Winfried W. C. Gieskes

Phytoplankton of the North Sea and its dynamics: A review

Abstract Phytoplankton is the major contributor to algal biomass and primary production of the North Sea, although crops of macroalgae can locally be up to 2000 g C.m−2 along the coast of the U.K. and Norway, and microphytobenthos dominates production in the shallow tidal flat areas bordering the coasts of England, the Netherlands, Germany and Denmark. Data collected since 1932 during the Continuous Plankton Recorder Survey show consistent patterns of geographical, seasonal and annual variation in the distribution of phytoplankton and its major taxonomic components. There is a trend of increased colouration in Recorder silks in the southern North Sea until approximately 1975 since when Colour levels (assumed to be indicative of algal biomass) have declined. In the eutrophic Dutch Wadden Sea the algal crop continued to increase; in Dutch coastal North Sea waters a trend of biomass increase reversed since 1984, apparently due to a reduction in Rhine river outflow. Long-term observations made at Helgoland since the 60s also show trends of increasing nutrients and phytoplankton biomass only to 1984. Adverse effects such as deoxygenation, foam formation and toxin production have been linked to mass concentrations of algae known as blooms. There is no evidence from existing reports for an increase in their frequency, although some years stand out with larger numbers. Occurrence of blooms can partly be explained by hydrographic conditions. More than 30 taxa are recognised as occurring in bloom proportions in the North Sea, approximately one third of which can be toxic. The crop of Bacillariophyceae (diatoms) is not likely to increase with eutrophication due to silicate limitation. An extensive subsurface maximum of armoured dinoflagellates, its abundance gouverned by hydrographic conditions, is the most characteristic feature of the central and northern North Sea in the summer months. Abundance, sometimes dominance, of picoplankton and of species that are not readily detected by microscopic observations has been documented by measurements of taxon-specific pigments such as chlorophyll b (green algae), alloxanthin (Cryptophyceae) and 19′ - hexanolyloxyfucoxanthin (Prymnesiophyceae or Haptophyceae). Analysis of time series of satellite images is a promising way to assess in a quantitative and, more important, synoptic way the patchy distribution of phytoplankton over large regions. Growth processes of the phytoplankton respond according to variables amenable to such satellite remote sensing. Empirical and theoretical relationships that can be established between chlorophyll a, 14C uptake, turbidity, stratification, suspended sediment type, irradiance and temperature in some well-investigated areas make remote sensing a potential tool to obtain reliable estimates of primary production in the whole North Sea. The 14C method for estimates of the rate of algal growth processes appears to agree reasonably well with other methods, both involving incubation of samples and in situ measurements of temporal changes of oxygen and pH. The level of net primary production is 250 g C.m−2.a−1 in the central North Sea, 150 to 200 g C.m−2.a−1 in the northern North Sea, and 200 g in the South. The main metabolic processes involved in phytoplankton growth have been modelled mathematically in terms of the most important controlling environmental parameters. Such parameters comprise not only those of a chemical signature (micro- and macronutrients, both inorganic and organic) but also physical effects of vertical mixing and sinking, and biological effects including allelopathic interactions, antibiotic excretions, vertical migration, and mortality due to grazing and parasitism. The balance between primary production and consumption of organic matter appears to vary both geographically and seasonally. The process of regeneration of primary products both in the water column and in and near the bottom seems to be of major importance. Future research should center around a study of growth-controlling parameters in laboratory culture experiments. The studies should include uptake of dissolved organic compounds by all taxonomic groups, including pico- and nanophytoplankton, and all aspects of ecosystem structure and function involving the relation between algae and microheterotrophs making up the small food web. There is a need to synthesize existing information on phytoplankton in the North Sea and the factors gouverning its growth, such as nutrients, river input and stratification intensity. Complicated inter-relationships and successional patterns between individual species which are limited by varying physiological requirements and adaptation to differing hydrographic regimes reemphasizes the importance of species identification in phytoplankton studies. Many future problems in phytoplankton research will not be resolved without accurate identification of algal species. Taxonomic expertise takes many years to acquire; there is at present a shortage of skills in this area and more resources should be turned towards training and long-term support.

Light-temperature interactions in the control of photosynthesis in Antarctic phytoplankton

SummaryDuring October/November 1983 photosynthetic responses of natural phytoplankton from the Scotia Sea and Bransfield strait to light and temperature were examined in incubators. Both assimilation numbers at saturating light levels and the slopes of the light-limited portions of the photosynthesis versus irradiance curves were smaller than in algae from lower latitudes. However, both parameters increased significantly with rising temperatures. Light-saturated photosynthesis on the average exhibited a Q10-value of ca. 4.2 between-1.5°C and +2°C. Light-limited photosynthesis between-1.5°C and +5°C rose at a rate corresponding to a Q10-value of roughly 2.6. Above +5°C, temperature enhancement of both light-saturated and light-limited photosynthetic rates was minimal or absent. Our results suggest that under extremely low temperatures light-limited photosynthetic rates become temperature-dependent due to changes in maximum quantum yields.

Biological control of short-term variations in the concentration of DMSP and DMS during a Phaeocystis spring bloom

Abstract In the spring of 1995, short-term variations in the concentration of particulate and dissolved dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) were monitored in the western Wadden Sea, a shallow coastal region in open connection with the North Sea. Significant correlations were found between abundance of Phaeocystis globosa and particulate DMSP; concentrations increased rapidly from 100 to 1650 n M in the middle of April. Highest DMS concentrations were found during the initial phase of the exponential growth of the bloom. DMS production and loss rates of DMSP and DMS were estimated experimentally during various phases of the bloom. DMS production and consumption were roughly in balance, with production only slightly exceeding consumption at the start of the bloom. Rates of production and consumption were highest during the exponential growth phase of Phaeocystis and declined in the course of the bloom (from 300–375 to less than 5 nmol dm −3 d −1 ). Demethylation of DMSP increased during the bloom (from 11 to 1300 nmol dm −3 d −1 ); it accounted for up to 100% of the DMSP loss at the end of the bloom. The shift from DMSP cleavage to demethylation in the course of a Phaeocystis bloom implies that DMS concentrations are not necessarily highest at the peak or towards the end of blooms.

The sensitivity of Emiliania huxleyi (Prymnesiophyceae) to ultraviolet‐b radiation

Emiliania huxleyi (Lohm.) Hay et Miller is an important component of the phytoplankton in open ocean waters. The sensitivity of this cosmopolitan alga to natural levels of UVB radiation has never been tested. Since DNA is believed to be a major target of natural UVB radiation (UVBR: 280–315 nm) in living cells, experiments with E. huxleyi were performed using growth rate reduction and DNA damage as indicators of UVBR stress. Specific growth rate, cell volume, pigment content, and CPD (cyclobutane pyrimidine dimer) formation (a measure for DNA damage) were followed during and after prolonged exposure of a series of cultures to a range of UVBR levels. E. huxleyi was found to be very sensitive to UVBR: at a daily weighted UVBR dose of only 400 J·m−2 ·d−1 (BEDDNA300nm), growth was halted. At this UVBR level, both cell volume and contents of the major photosynthetic and photoprotective pigments had increased. The UVBR vulnerability of E. huxleyi cannot be explained by a high potential for cyclobutane thymine dimer formation (the most abundant CPD type) due to a high T content of nuclear DNA: the CG content of this E. huxleyi strain is high (68%) compared with other species. The high UVBR sensitivity may be related to the stage of the cell cycle during UVBR exposure, in combination with low repair capacity. It is concluded that E. huxleyi may experience UVBR stress through the formation of cyclobutane pyrimidine dimers, with subsequent low repair capacity and thereby arrest of the cell cycle.

Temperature, light, and the dimethylsulfoniopropionate (DMSP) content of Emiliania huxleyi (Prymnesiophyceae)

The precursor of the volatile S-compound dimethylsulfide (DMS), dimethylsulfoniopropionate (DMSP), is produced by marine microalgae, notably by Prymnesiophyceae. The production of DMSP by an axenic isolate of Emiliania huxleyi (Lohm.) Hay et Mohler under different temperature and light conditions was studied as a first step towards understanding the role of DMSP-producing algae in climate regulation. Both light and temperature affected growth rate and cell size in batch cultures, but the concentration of DMSP in the cells was dependent on temperature only: at low temperature DMSP accumulated. This physiological response, assumed to be characteristic of DMSP-producing microalgae in general, is in line with the correlation that has been found between elevated concentrations of the DMS oxidation product MSA in ice core slices corresponding with low sea surface temperatures. Apparently, DMS does not play the role in climate regulation formulated in the CLAW hypothesis that proposes a feedback mechanism in which elevated temperatures lead to an increase in albedo via DMS-derived cloud condensation nuclei.

GROWTH AND FLUORESCENCE CHARACTERISTICS OF ULTRAPLANKTON ON A NORTH SOUTH TRANSECT IN THE EASTERN NORTH-ATLANTIC

In the summer of 1989 vertical profiles of chlorophyll a were taken in the North Atlantic. Stations were located along a transect following longitude 20°W, between 60 and 33°N. Maximum chlorophyll a levels were located near the surface in the north (2 μl−1 but became gradually deeper towards the south (0.3 μ l−1 at a depth of 80–110m). Primary production varied only by a factor of 2.3 (226–533 mg C m−2 day−1), whereas the assimilation ratio showed far less variation (15.46±4.61 mg C mg Chl−1 day−1, n = 14). The northern part of the transect showed no thermal stratification, and nutrients were plentiful. Phytoplankton was distributed homogeneously, with a nearly constant signature of the cellular fluorescence characteristics as monitored by flow-cytometry. The dominant taxa were small prymnesiophytes (Emiliania huxleyi) and chroococcoid phycoerythrin-containing cyanobacteria. Cells were healthy, resulting in net growth rates (growth rate minus grazing) of up to 0.75 divisions per day. Of the total primary production, 68% was actually converted into new plant carbon. Midway along the transect (47°N) there was a sharp thermocline at a depth of 35 m. All nutrients (nitrogen, phosphate and silicate) were depleted in the upper water layer. Cyanobacteria dominated the phytoplankton community in numbers varying from 12,000 to 39,000 cells ml−1. Stabilization of the water column was apparently in progress, as could be derived from the gradual increase in mean cellular red fluorescence of the cyanobacteria with depth. Furthermore, net growth rates of the algae were relatively low, as was the proportion of primary production resulting in an actual increase of plant carbon (42%). At the southernmost station (33°N), the situation was typical of the oligotrophic subtropics: a sharp thermocline at 35 m, the 0.1% surface incident irradiance level at 150 m, and a nitracline at a depth of 100–120m. The most prominent (pico)plankters present were prochlorophytes, predominantly at the deep chlorophyll maximum (80–100 m, up to 95,000 cells ml−1). Cyanobacteria co-occurred but only in small numbers (maximum of 5500 ml−1), mainly in surface waters. The vertical (physical) stability of the water column was pronounced, and the mean cellular red fluorescence signal of cells found at the bottom of the euphotic zone was consequently higher than in surface waters. Most remarkable were the net growth rates of the various species suggesting a highly dynamic phytoplankton community. Extreme growth rates included positive as well as negative values which were not always consistent with the greatest abundance of certain species. Despite these rapid changes, the actual increase in plant carbon was only 14% of the primary production.

Relaxed eddy accumulation measurements of the sea-to-air transfer of dimethylsulfide over the northeastern Pacific

[1] Gas transfer rates were determined from relaxed eddy accumulation ( REA) measurements of the flux of dimethylsulfide (DMS) over the northeastern Pacific Ocean. This first application of the REA technique for the measurement of DMS fluxes over the open ocean produced estimates of the gas transfer rate that are on average higher than those calculated from commonly used parameterizations. The relationship between the total gas transfer rate and wind speed was found to be gas k(gas) = 0.53 (+/-0.05) U-10(2). Because of the effect of the airside resistance, the waterside transfer rate was up to 16% higher than kgas. Removal of the airside transfer component from the total transfer rate resulted in a relation between wind speed and waterside transfer of k(660) = 0.61 (+/-0.06) U-10(2). However, DMS fluxes showed a high degree of scatter that could not readily be accounted for by wind speed and atmospheric stability. It has to be concluded that these measurements do not permit an accurate parameterization of gas transfer as a function of wind speed.

Salinity-dependent diatom biosilicification implies an important role of external ionic strength

The role of external ionic strength in diatom biosilica formation was assessed by monitoring the nanostructural changes in the biosilica of the two marine diatom species Thalassiosira punctigera and Thalassiosira weissflogii that was obtained from cultures grown at two distinct salinities. Using physicochemical methods, we found that at lower salinity the specific surface area, the fractal dimensions, and the size of mesopores present in the biosilica decreased. Diatom biosilica appears to be denser at the lower salinity that was applied. This phenomenon can be explained by assuming aggregation of smaller coalescing silica particles inside the silica deposition vesicle, which would be in line with principles in silica chemistry. Apparently, external ionic strength has an important effect on diatom biosilica formation, making it tempting to propose that uptake of silicic acid and other external ions may take place simultaneously. Uptake and transport of reactants in the proximity of the expanding silica deposition vesicle, by (macro)pinocytosis, are more likely than intracellular stabilization and transport of silica precursors at the high concentrations that are necessary for the formation of the siliceous frustule components.

Phaeocystis globosa (Prymnesiophyceae) colonies: hollow structures built with small amounts of polysaccharides

In both field samples and cultures the total amount of sugar and of carbon in colonies of Phaeocystis globosa was correlated with colony surface area, suggesting a hollow structure. A conceptual model based on biochemical data and on the assumption that the mucus occurs as a layer of a fixed thickness, irrespective of colony size, predicts that the thickness of the layer is 7 μm. A confocal laser scanning microscope image of fluorescently labelled mucus confirmed this view of the colony structure. The measured contents of carbon and sugar per cell (including mucus) were constant for all colony sizes. Cells in laboratory cultures contained 122 pg C, which is twice the value for cells obtained at a field station in the North Sea (57 pg). In contrast, sugar per cell was higher in the field than in exponentially growing cultures. Therefore the percentage of sugar carbon relative to total carbon of colonies in the field was higher (19–35%) than that in cultures (10%). Highest values were found at low ambient n...

Nanoscale uniformity of pore architecture in diatomaceous silica: a combined small and wide angle x‐ray scattering study

Combined small and wide angle X‐ray scattering (SAXS and WAXS) analysis was applied to purified biogenic silica of cultured diatom frustules and of natural populations sampled on marine tidal flats. The overall WAXS patterns did not reveal crystalline phases (WAXS domain between 0.07 to 0.5 nm) in this biogenic silica, which is in line with previous reports on the amorphous character of the SiO2 matrix of diatom frustules. One exception was the silica of the pennate species Cylindrotheca fusiformis Reimann et Lewin, which revealed wide peaks in the WAXS spectra. These peaks either indicate the presence of a yet unknown crystalline phase with a repetitive distance (d‐value ≈0.06 nm) or are caused by the ordering of the fibrous silica fragments; numerous girdle bands. The SAXS spectra revealed the size range of pores (diameter d between 3.0 and 65 nm), the presence of distinct pores (slope transitions), and structure factors (oscillation of the spectra). All slopes varied in the range of −4.0 to −2.5, with two clear common regions among species: d < 10 nm (slopes –4, denoted as region I and also called the Porod region), and 10.0 < d < 40.0 nm (slopes −2.9 to −3.8, denoted as region II). The existence of these common regions suggests the presence of comparable form (region I) and structure (region II) factors, respectively the shape of the primary building units of the silica and the geometry of the pores. Contrast variation experiments using dibromomethane to fill pores in the SiO2 matrix showed that scattering was caused by pores rather than silica particles. Electron microscopic analysis confirmed the presence of circular, elliptical, and rectangular pores ranging in size from 3 to 65 nm, determining the structure factor. The fine architecture (length/width ratio of pore diameters) and distribution of the pores, however, seemed to be influenced by environmental factors, such as the salinity of and additions of AlCl3 to the growth medium. The results indicate that diatoms deposit silica with pores <50 nm in size and are highly homologous with respect to geometry. Consequently, it is suggested that in diatoms, whether pennate or centric, the formation of silica at a nanoscale level is a uniform process.