Dinoflagellate cysts in recent marine sediments of the western coast of the Bering Sea

The quantitative and qualitative composition of live dinoflagellate cysts was studied in the upper two-centimeter layer of recent marine sediments that were collected at 19 stations in the coastal waters of the western Bering Sea. A total of 28 types of identified cysts belonged to the following 11 genera: Alexandrium, Diplopsalis, Ensiculifera, Gonyaulax, Gyrodinium, Pentapharsodinium, Polykrikos, Preperidinium, Protocera- tium, Protoperidinium, and Scrippsiella. The morphology of dinoflagellate cysts from recent sediments of Russian seas, such as the shape, the size, and also the structure of the phragma, including the processes and the archeopyle, was described comprehensively for the first time. Cysts of the species Gonyaulax spinifera, Pentapharsodinium dalei, Protoceratium reticulatum, Protoperidinium americanum, P. conicoides, P. subinerme, Scrippsiella crystallina, and S. trochoidea were the most widespread. Those of the potentially toxic species Alexandrium tamarense were also widely distributed and prevailed in the studied area. Their concentration varied from 0 to 25 860 cells/cm³; the maximum concentration was recorded in Pavel Bay, Koryak Okrug, and Kamchatka.     

Subsurface phytoplankton blooms fuel pelagic production in the North Sea

The seasonal phytoplankton biomass distribution pattern in stratifiedtemperate marine waters is traditionally depicted as consistingof spring and autumn blooms. The energy source supporting pelagicsummer production is believed to be the spring bloom. However,the spring bloom disappears relatively quickly from the watercolumn and a large proportion of the material sedimenting tothe bottom following the spring bloom is often comprised ofintact phytoplankton cells. Thus, it is easy to argue that thespring bloom is fueling the energy demands of the benthos, butmore difficult to argue convincingly that energy fixed duringthe spring bloom is fueling the pelagic production occurringduring summer months. We argue here that periodic phytoplanktonblooms are occurring during the summer in the North Sea at depthsof >25 m and that the accumulated new production [sensu (Dugdaleand Goering, Limnol. Oceanogr., 12, 196–206, 1967)] occurringin these blooms may be greater than that occurring in the springbloom in the same regions. Thus, such blooms may explain apparentdiscrepancies in production yields between different temperatemarine systems.     

Abundance and distribution of toxic Alexandrium tamarense resting cysts in the sediments of the Chukchi Sea and the eastern Bering Sea

Abundance and distribution of the toxic dinoflagellate Alexandrium tamarense species complex resting cyst were investigated in the eastern Bering Sea and the Chukchi Sea for the first time. Sediment samples (top 0–3 cm depth) were collected from the continental shelf of the eastern Bering Sea (17 stations) and the Chukchi Sea (13 stations) together with a long core sample (top 0–21 cm depth) from one station in the Chukchi Sea during 2009–2012. The cysts were enumerated using the primuline staining method. Species identification of the cysts was carried out with multiplex PCR assay and the plate morphology of vegetative cells germinated from cysts in the both areas. Alexandrium cysts were widely detected in the both areas, ranging from not detected (<1 cysts cm⁻³) to 835 cysts cm⁻³ wet sediment in the eastern Bering Sea and from not detected (<1 cysts cm⁻³) to 10,600 cysts cm⁻³ in the Chukchi Sea, and all isolated cysts were genetically and morphologically identified as the North American clade A. tamarense. Their cysts were mainly distributed in the shallow continental shelf where the water depth was less than 100 m in both areas. The cysts were detected from the deep layer (18–21 cm depth of sediment core) of the long core sample. The present study confirmed the abundant existence of A. tamarense with wide range of distribution in these areas. This fact suggests that A. tamarense vegetative cells have appeared in the water column in the both areas. Furthermore, these abundant cyst depositions indicate that this species originally distributed in the Arctic and subarctic regions and well adapted to the environments in the marginal ice zone.     

Dinoflagellate cysts as indicators of water quality and productivity in British Columbia estuarine environments

Palynological analyses of 60 surface sediment samples from estuarine environments near Vancouver Island, including the Georgia Strait (GS), the Effingham (EFF) and the Seymour–Belize (SB) Inlets were performed in order to document the distribution of dinoflagellate cyst assemblages and their relationship to hydrographic conditions, productivity and nutrient concentrations. We tested transfer functions using the analogue method, and suggest that dinoflagellate cyst assemblages can be used to reconstruct primary productivity, temperature and salinity. The EFF and SB Inlets are characterized by a dominance of autotrophic taxa, particularly Operculodinium centrocarpum, whereas the Protoperidinioid and gymnodinial heterotrophic taxa such as Quinquecuspis concreta and Brigantedinium spp. dominate the assemblages of the GS. Multivariate analysis shows that this distribution is closely linked to primary productivity, sea-surface temperature (SST) and spring silica concentration. The abundance of autotrophic taxa in the EFF and SB Inlets is associated with high primary productivity and low summer SST, indicating summer upwelling of coastal British Columbia, whereas the heterotrophic taxa that characterizes the GS assemblages are related to low productivity, high summer SST and high silica concentration during spring. Multivariate analysis shows that the most important environmental parameters related to dinocyst distribution in the restricted embayment of the GS, are distance to the shore, distance to Vancouver Harbor, spring sea surface salinity (SSS), spring phosphate concentration and spring productivity. The autotrophic taxa are generally more common in coastal and shallow waters, but Spiniferites ramosus and Pentapharsodinium dalei show an opposite correlation to spring productivity and salinity. P. dalei is particularly abundant around Vancouver Harbor, near highly urbanized shores and within the Fraser River plume, where salinity is low and spring productivity and continental runoff are high. S. ramosus shows its highest abundance on the western coast of GS. Protoperidinioid and gymnodinial cysts characterize distal zones within the central and southern GS that are associated with a mixture of brackish waters coming from the Fraser River and deep upwelling waters entering the GS via Juan de Fuca Strait. The relationship between dinoflagellate cyst assemblages and primary productivity in these estuarine systems differs from that in oceanic and outer neritic zones, where the abundance of heterotrophic taxa is commonly associated with upwelling and high productivity.     

A plea for more ecology in phytoplankton ecology

When looking for a pattern of phytoplankton behaviour across trophic gradients, we need to cross the boundaries between different disciplinary areas, from autoecology to systems ecology, because eutrophication is a complex process which involves different time scales and different levels of community structure. Thus, we submit our observations to the muddled conceptual world of assemblage ecology. These inaccuracies arise, for example, from both species and community arguments; eutrophication as a fertilization or a metabolic phenomenon; and the notions frequently interwoven of pattern, process and rules. We suggest that it is advantageous to tackle this issue from the perspective of general ecology, rather than from a specifically planktonic orientation. In this way, useful general ecological tools, for example, time series and assembly-rule studies, can be used. Time-series study allows the dynamics of any variable to be described or to show that long term variable fluctuations may sometimes be unregulated, in response to some exogenous factor. Rules of assembly help us to resolve which traits are selectively involved during the eutrophication process. In this context, we advocate (1) the use of traits instead of morphospecies in phytoplankton studies, (2) looking for the dynamic patterns of phytoplankton with eutrophication, (3) the use of time series techniques to study phytoplankton trajectories, (4) the use of assembly rules to discern patterns in the formation of multispecies assemblages, (5) the consideration of the pelagic food-web in studies of phytoplankton dynamics and, as an overall suggestion, to borrow knowledge and inspiration from general ecology.     

The strain concept in phytoplankton ecology

Laboratory cultures are important tools for investigating the biology of microalgae, allowing experimentation under controlled conditions. This control is critical for comparative studies, such as those often used to investigate intra-specific variation in properties of interest. By holding the environment constant, the experimentalist can gain insight into the genetic basis of phytoplankton phenotypes and by extension, into the adaptive history of those genotypes. In most cases the adaptations of interest are those that the algae have evolved in response to their natural environment. However, here it is argued that such experiments may instead reveal evolutionary adaptations to, and/or non-adaptive changes induced by, the culture conditions under which the alga is maintained. We present a review of the processes of evolution as they pertain to microalgal culture, and illustrate this discussion with examples of in-culture evolution from both within and outside the field of phycology. With these considerations in mind, recommendations are made for experimental practice focusing on comparative physiology, for which the effects of in-culture evolution are particularly confounding. Finally we argue that, although problematic in some contexts, the evolutionary propensities of phytoplankton cultures actually present an important opportunity for experimental evolutionary research with direct environmental significance.     

Contrasting nitrogen uptake by diatom and Phaeocystis-dominated phytoplankton assemblages in the North Sea

This paper documents ambient concentrations of nutrients in the Belgian coastal waters of the North Sea during the spring of 1996 and 1997. The paper elaborates the differences of uptake rates of oxidised nitrogen (NO₃⁻) and reduced nitrogen (NH₄ and urea) by Phaeocystis and diatoms. The nitrogen concentrations were dominated by NO₃⁻ with a maximum concentration of 30 μM (January 1997) and 40 μM (March 1996). In 1996, Phaeocystis dominated the spring biomass with a maximum of 521 μg C l⁻¹, while maximum diatom biomass was 174 μg C l⁻¹. In 1997, the maximum Phaeocystis spring biomass was 1600 μg C l⁻¹ and diatom maximum biomass was below 100 μg C l⁻¹. A maximum bacteria biomass of about 55 μg C l⁻¹ was observed in mid-May 1996. The maximum nitrogen uptake rates were recorded during spring and were dominated by NO₃⁻ (0.005 h⁻¹ in 1996 and 0.032 h⁻¹ in 1997). Maximum specific NH₄ uptake rates were between 0.005 h⁻¹ in May 1996 and 0.006 h⁻¹ in April 1997. The NO₃⁻ uptake rates displayed exponential decrease versus increasing ambient reduced nitrogen concentrations (ammonium and urea), whereas the reduced nitrogen uptake increased but never compensated the decreased nitrate uptake. The NH₄ uptake kinetics of diatoms displayed lower v_max compared to Phaeocystis. Consequently, Phaeocystis showed ability to increase their NH₄ uptake capacity when more NH₄ became available while diatoms failed to do so, after ammonium had exceeded their saturation concentration (>1 μM). Although reduced nitrogen has a negative effect on the uptake of NO₃⁻, Phaeocystis have more advantage than diatoms on the uptake of ammonium. This might be contributing to the biomass domination shown by Phaeocystis over extended periods in spring.     

The warming trend at Helgoland Roads,North Sea: phytoplankton response

We combine the temperature and phytoplankton data from one of the longest aquatic data sets in history, the Helgoland Roads (North Sea, 54°11.3N, 7°54.0E) timeseries to evaluate the effects of climate change on the base of marine food webs. The data shows that, despite an obvious warming of 1.1°C since 1962, the mean diatom day of the algal spring bloom is delayed and shifted to the end of the first quarter of the year. This is apparently related to a warming of the autumn (October–December) temperatures. It is the first indication of a warming related shift in phytoplankton succession, the consequences of which would range from lifecycle/food resource mismatches to regime shifts in the North Sea system.     

Diet and fish stock availability as possible factors in the mass death of auks in the North Sea

In February 1983 more than 30 000 birds died on the North Sea coasts of England and Scotland. The birds were unoiled, and there was no immediately obvious cause of death. Stomach contents of auks killed in this incident were examined to ascertain whether they provided any indication of the causal factors. Analysis of food remains showed differences between sample sites and between guillemots, Una aalge (Pont.), and razorbills, Alca tarda L. In all areas gadoid remains comprised a higher, and sandeels a lesser, proportion of the stomach contents in guillemots than in razorbills. The relative importance of clupeid remains varied with area. These differences, combined with current trends in the fisheries biology of major prey species, provided an ecological basis for a hypothesis explaining the incident. It was concluded that starvation was probably a major factor in the death of the auks, and that this may have been related to a combination of adverse weather conditions and changing patterns of abundance in North Sea sprat, Sprattus sprattus (L.), populations.     

Factors regulating germination of resting cysts of the spring bloom dinoflagellate Scrippsiella hangoei from the northern Baltic Sea

The role of cyst germination as a factor in bloom initiationwas investigated for the dinoflagellate Scrippsiella hangoei(Schiller) Larsen from the northern Baltic Sea. This speciesblooms in very cold, often ice-covered waters, and is responsiblefor a significant fraction of the production in that region.Dormancy, temperature, oxygen and light were studied as factorspotentially controlling the germination of S.hangoei restingcysts. Laboratory-stored and field-collected cysts began togerminate in December following a mandatory dormancy periodlasting 6 months. Germination after this maturation intervalwas maximal when temperatures were within a 0–9°C‘window’. Mandatory dormancy is therefore the primaryfactor regulating the timing of germination in this species,as temperatures in the natural environment normally fall withinthis range at the time when S.hangoei cysts deposited the precedingyear have matured. Non-optimal temperatures, darkness and lowoxygen conditions all maintain a state of quiescence in thecysts. Cysts could germinate in darkness, but the rate of excystmentwas significantly higher in the light. Likewise, excystmentwas completely inhibited in anoxic conditions and was reducedunder severe hypoxia, with normal germination under moderatehypoxic concentrations. Temporary exposure to high sulfide concentrationspermanently reduced germination potential, indicating that S.hangoeicysts have low resistance to oxygen deficiency. Prolonged periodsof anoxia at the sediment surface, as frequently occurs in thestudy area, might reduce the size of the viable cyst pool andthus, alter the magnitude of the inoculum for S.hangoei bloominitiation. Together, these internal and external regulatoryfactors play important roles in the bloom dynamics of this importantdinoflagellate.     

Equilibrium/steady-state concept in phytoplankton ecology

This paper summarises the outcomes of the 13^th Workshop of the International Association of Phytoplankton Taxonomy and Ecology (IAP). The authors mostly addressed their contributions on the following topics: the effect of trophic state on the attainment of a steady-state; the establishment of equilibria in deep and shallow lakes; the role of spatial heterogeneity, disturbance, and stress in the establishment of equilibrium assemblages; the mechanisms leading to the steady state; the frequency and longevity of equilibrium phases, and the role of morphological and physiological plasticity of phytoplankton in maintaining the (apparently) same populations under different environmental conditions. The composition of steady-state assemblages is compared to that of phytoplankton functional groups (coda). Those functional associations recognised as steady-state assemblages appear to be strongly K-determined in many instances.     

Summer phytoplankton distributions in the Weddell Sea

Summary The quantitative composition of phytoplankton was studied along a transect of 14 hydrographic stations, between the southern coast of the Weddell Sea and the Antarctic Peninsula, during the austral summer of 1984–1985. The most apparent feature of the phytoplankton distribution was the presence of a bloom of Phaeocystis at a frontal zone over the shelf break, and the marked contrast between a southern region, with high phytoplankton biomass, and a poorer region north of the shelf break. The most widely distributed diatom genus was Nitzschia (Fragilariopsis section). The phytoplankton assemblage of the southern region included the silicoflagellate Distephanus speculum, the diatom Rhizosolenia alata and several heterotrophic dinoflagellates such as Protoperidinium antarcticum and P. applanatum. The northern assemblage could be characterized by the abundance of flagellates and small dinoflagellates, and by diatoms such as Chaetoceros criophilum, Corethron criophilum Nitzschia kerguelensis and other Nitzschia species of the Fragilariopsis section.     

Physiological diversity of Roseobacter clade bacteria co-occurring during a phytoplankton bloom in the North Sea

Organisms of the Roseobacter clade are an important component in marine ecosystems, partially due to their metabolic variety. Not much is known, however, about the physiological diversity of different roseobacters present within one habitat. By using serial dilution cultures with low-nutrient media seven roseobacter strains, co-occurring during a phytoplankton bloom in the southern North Sea, were obtained in this study. Physiological characterization exhibited distinct substrate spectra of the isolates. Although no isolate showed growth on algal osmolyte dimethylsulfoniopropionate (DMSP), feeding experiments revealed that all new strains converted [²H₆]DMSP into a variety of volatile compounds. Six strains mainly decomposed DMSP via the demethylation pathway, but four strains were also capable of cleaving DMSP to DMS and acrylate. It is hypothesized that the great physiological diversity of the roseobacters reflects their ability to inhabit different ecological niches and enables the organisms to cope differently with changing substrate supplies during phytoplankton blooms. Denaturing gradient gel electrophoresis and sequencing of excised bands resulted in detection of five additional roseobacters. Three of these sequences showed affiliation with three of the four major clusters of the Roseobacter clade, consisting predominantly of uncultured organisms (i.e. the Roseobacter clade-affiliated (RCA)), the NAC11-7 and the CHAB-I-5 clusters.     

Recent Dinoflagellate cyst distribution in the North Canary Basin,NW Africa

The North Canary Basin (NW Africa) falls within a major eastern boundary upwelling system. This part of the coastal upwelling system is seasonal and is characterised by the development of large filaments migrating seawards. Hence, 16 samples from this location were selected to identify an “upwelling signal” in the composition of the dinoflagellate cyst assemblages.

Samples closest to the most intense upwelling cells are dominated by L. machaerophorum and G. catenatum and Protoperidinium spp. These make up the “upwelling signal” characteristic for the system. Moreover, the “upwelling signal” can be advected offshore, with filaments that may extend as far as 300 km. Finally, the finding of cysts from G. catenatum, a toxic dinoflagellate, raises the need for a better understanding of the relationship between its presence and distribution in the region, and the coastal upwelling system.     

Dynamics in bacterial surface properties of a natural bacterial community in the coastal North Sea during a spring phytoplankton bloom

The hydrophilic and hydrophobic properties of single cells of natural bacterioplankton communities were determined using a recently developed staining method combined with confocal laser scanning microscopy and advanced image analysis. On an average, about 50% of the bacterial cell area was covered by hydrophobic and only 16% by hydrophilic properties, while about 72% was covered by the genome. However, the size of these properties was independent of the bacterial cell size. Bacterial hydrophobicity was positively correlated with ambient NH(4)(+) concentrations and negatively correlated with overall bacterial abundance. The expression of hydrophilicity was more dynamic. Over the spring phytoplankton bloom, the bacterioplankton ratio(phil/phob) repeatedly reached highest values shortly before peaks in bacterioplankton abundance were observed, indicating a direct and fast response of bacterial surface properties, especially hydrophilicity, to changing environmental conditions. Compared to bacterial strains, recently studied with the same method, cells of marine bacterioplankton communities are much smaller and less frequently covered by hydrophobic or hydrophilic properties. While the percentage area covered by the genome is essentially the same, the percentage area covered by hydrophobic or hydrophilic properties is much smaller.