Decadal basin-scale changes in diatoms, dinoflagellates, and phytoplankton color across the North Atlantic

The Continuous Plankton Recorder (CPR) survey has been used to characterize phytoplankton and zooplankton space-time dynamics in the North Sea since 1931 and in the North Atlantic since 1939. Phytoplankton biomass is assessed from these samples by visual assessment of the green color of the silk mesh, the Phytoplankton Color Index (PCI), and the total count of diatoms and dinoflagellates. Species with a frequency of occurrence greater than 1% in the samples are used as indicator species of the community. We investigated (1) long-term fluctuations of phytoplankton biomass, total diatoms, and total dinoflagellates; (2) geographical variation of patterns; (3) the relationship between phytoplankton and climate forcing in the North Atlantic CPR samples; (4) the relative contribution of diatoms and dinoflagellates to the PCI; and (5) the fluctuations of the dominant species over the period of survey to provide more information on the processes linking climate to changes in the phytoplankton community. As a result of the differences in microscopic analysis methods prior to 1958, our analyses were conducted for the period ranging from 1958 to 2002. The North Atlantic was divided into six regions identified through bathymetric criteria and separated along a North-South axis. Based on 12 monthly time series, we demonstrate increasing trends in PCI and total dinoflagellates and a decrease in total diatoms.

Differential contribution of diatoms and dinoflagellates to phytoplankton biomass in the NE Atlantic Ocean and the North Sea

Sampling by the continuous plankton recorder (CPR) survey over the North Atlantic Ocean and the North Sea has enabled long-term studies of phytoplankton biomass. Analysis of an index of phytoplankton biomass, the phytoplankton colour index (PCI), has previously shown an increase in phytoplankton biomass in the NE Atlantic. In the current study, further investigations were conducted to determine the contribution of diatom and dinoflagellate cell counts to the PCI, their fluctuations over the last 45 yr and their geographical variations in the eastern North Atlantic and the North Sea. An increased contribution of dinoflagellates to the PCI was revealed over the south NE Atlantic and the northern North Sea. In contrast, the contribution of diatoms decreased in the north NE Atlantic and the northern North Sea. No discernible trends were found in the other regions of the North Sea. The relative contributions of diatoms and dinoflagellates to the PCI led to the identification of 3 geographically distinct dynamic regimes in the diatom/dinoflagellate dynamics in the NE Atlantic and the North Sea. Finally, it is stressed that the discrepancy observed in the patterns of PCI and diatom and dinoflagellate cell counts suggests that changes in PCI do not reflect changes in the community structure and that the exclusive use of PCI is not adequate to investigate the long-term trends in the trophic link between phytoplankton and herbivorous zooplankton.

Automatic classification of field-collected dinoflagellates by artificial neural network

Automatic taxonomic categorisation of 23 species of dinoflagellates was demonstrated using field-collected specimens. These dinoflagellates have been responsible for the majority of toxic and noxious phytoplankton blooms which have occurred in the coastal waters of the European Union in recent years and make severe impact on the aquaculture industry. The performance by human 'expert' ecologists/taxonomists in identifying these species was compared to that achieved by 2 artificial neural network classifiers (multilayer perceptron and radial basis function networks) and 2 other statistical techniques, k-Nearest Neighbour and Quadratic Discriminant Analysis. The neural network classifiers outperform the classical statistical techniques. Over extended trials, the human experts averaged 85% while the radial basis network achieved a best performance of 83%, the multilayer perceptron 66%, k-Nearest Neighbour 60%, and the Quadratic Discriminant Analysis 56%.

Modelling the Stoichiometric Regulation of C-Rich Toxins in Marine Dinoflagellates

Toxin production in marine microalgae was previously shown to be tightly coupled with cellular stoichiometry. The highest values of cellular toxin are in fact mainly associated with a high carbon to nutrient cellular ratio. In particular, the cellular accumulation of C-rich toxins (i.e., with C:N > 6.6) can be stimulated by both N and P deficiency. Dinoflagellates are the main producers of C-rich toxins and may represent a serious threat for human health and the marine ecosystem. As such, the development of a numerical model able to predict how toxin production is stimulated by nutrient supply/deficiency is of primary utility for both scientific and management purposes. In this work we have developed a mechanistic model describing the stoichiometric regulation of C-rich toxins in marine dinoflagellates. To this purpose, a new formulation describing toxin production and fate was embedded in the European Regional Seas Ecosystem Model (ERSEM), here simplified to describe a monospecific batch culture. Toxin production was assumed to be composed by two distinct additive terms; the first is a constant fraction of algal production and is assumed to take place at any physiological conditions. The second term is assumed to be dependent on algal biomass and to be stimulated by internal nutrient deficiency. By using these assumptions, the model reproduced the concentrations and temporal evolution of toxins observed in cultures of Ostreopsis cf. ovata, a benthic/epiphytic dinoflagellate producing C-rich toxins named ovatoxins. The analysis of simulations and their comparison with experimental data provided a conceptual model linking toxin production and nutritional status in this species. The model was also qualitatively validated by using independent literature data, and the results indicate that our formulation can be also used to simulate toxin dynamics in other dinoflagellates. Our model represents an important step towards the simulation and prediction of marine algal toxicity.

Increased blooms of a dinoflagellate in the NW Atlantic

Sampling by the Continuous Plankton Recorder (CPR) over the NW Atlantic from 1960 to 2000 has enabled long-term studies of the larger components of the phytoplankton community, highlighting various changes, particularly during the 1990s. Analysis of an index of phytoplankton biomass, the Phytoplankton Colour Index (PCI) has revealed an increase over the past decade, most marked during the winter (December to February) months. Examination of the structure of the community using multiple linear-regression models indicates that the winter phytoplankton community composition has changed markedly in the 1990s compared to the 1960s. One phytoplankter, the dinoflagellate Ceratium arcticum (Cleve), has undergone dramatic changes in abundance during this period, with pronounced large winter blooms and decreased autumnal levels, and its contribution to the Phytoplankton Colour index values has increased significantly. Other dominant species in the phytoplankton community, both diatoms and dinoflagellates, did not show the same variations over the examined time period. It is suggested that the response of C. arcticum is probably a result of previously reported changes in stratification in the NW Atlantic, due to dynamic hydro-climatic (freshening and cooling) events.

A Possible Relationship Between Chitinozoa And Tintinnids

The biological affinity of the extinct microfossil order chitinozoa has been the source of much discussion in the fifty years since they were first discovered. Within this period these flask-shaped, organic-walled organisms have been variously attributed to rhizopods, flagellates, tintinnids, chrysomonads, metazoan eggs, dinoflagellates, and fungi. Most of these suggested relationships were made before it was recognised that chitinozoans were encapsulated and must therefore be resting cysts or eggs and not active individuals. There are no living organisms which combine all the characteristics of the chitinozoa. Of all the possibilities, a grouping of flask-shaped cysts which have been found in present-day marine plankton and sediment comes closest to characterising the morphology of chitinozoa. This grouping of flask-shaped cysts includes forms which have been found within tintinnid loricae. Another modern cyst type Pacillina arctica, which is believed to be a ciliate cyst, comes close to replicating the morphology of the chitinozoan genus Hoegisphaera. This paper discusses the structure of tintinnid, other flask-shaped cysts and Pacillina arctica in relation to chitinozoan morphology, drawing attention to similarities and differences. The occurrence and distribution of these cyst forms in present-day plankton is also described and interpreted.

Dynamics of phytoplankton communities during late summer around the tip of the Antarctic Peninsula

The composition and distribution of phytoplankton assemblages around the tip of the Antarctic Peninsula were studied during two summer cruises (February/March 2008 and 2009). Water samples were collected for HPLC/CHEMTAX pigment and microscopic analysis. A great spatial variability in chlorophyll a (Chl a) was observed in the study area: highest levels in the vicinity of the James Ross Island (exceeding 7 mg m−3 in 2009), intermediate values (0.5 to 2 mg m−3) in the Bransfield Strait, and low concentrations in the Weddell Sea and Drake Passage (below 0.5 mg m−3). Phytoplankton assemblages were generally dominated by diatoms, especially at coastal stations with high Chl a concentration, where diatom contribution was above 90% of total Chl a. Nanoflagellates, such as cryptophytes and/or Phaeocystis antarctica, replaced diatoms in open-ocean areas (e.g., Weddell Sea). Many species of peridinin-lacking autotrophic dinoflagellates (e.g., Gymnodinium spp.) were also important to total Chl a biomass at well-stratified stations of Bransfield Strait. Generally, water column structure was the most important environmental factor determining phytoplankton communities’ biomass and distribution. The HPLC pigment data also allowed the assessment of different physiological responses of phytoplankton to ambient light variation. The present study provides new insights about the dynamics of phytoplankton in an undersampled region of the Southern Ocean highly susceptible to global climate change.

Nitrogen uptake of phytoplankton assemblages under contrasting upwelling and downwelling conditions: The Ría de Vigo, NW Iberia

Regenerated production (including organic nitrogen) is shown here to be important in the Ria de Vigo (Galicia, NW Iberia) in supporting both harmful algal bloom communities during the downwelling season, but also (to a lesser extent) diatom communities during stratified periods of weak to moderate upwelling. The Galician Rias, situated in the Iberian upwelling system, are regularly affected by blooms of toxic dinoflagellates, which pose serious threats to the local mussel farming industry. These tend to occur towards the end of summer, during the transition from upwelling to downwelling favourable seasons, when cold bottom shelf waters in the rias are replaced by warm surface shelf waters. Nitrate, ammonium and urea uptake rates were measured in the Ria de Vigo during a downwelling event in September 2006 and during an upwelling event in June 2007. In September the ria was well mixed, with a downwelling front observed towards the middle of the ria and relatively high nutrient concentrations (1.0-2.6 mu mol L-1 nitrate; 1.0-5.6 mu mol L-1 ammonium; 0.1-0.8 mu mol L-1 phosphate; 2.0-9.0 mu mol L-1 silicic acid) were present throughout the water column. Ammonium represented more than 80% of the nitrogenous nutrients, and the phytoplankton assemblage was dominated by dinoflagellates and small flagellates. In June the water column was stratified, with nutrient-rich, upwelled water below the thermocline and warm, nutrient-depleted water in the surface. At this time, nitrate represented more than 80% of the nitrogenous nutrients, and a mixed diatom assemblage was present. Primary phytoplankton production during both events was mainly sustained by regenerated nitrogen, with ammonium uptake rates of 0.035-0.063 mu mol N L-1 h(-1) in September and 0.078-0.188 mu mol N L-1 h(-1) in June. Although f-ratios were generally low (<0.2) in both June and September, a maximum of 0.61 was reached in June due to higher nitrate uptake (0.225 mu mol N L-1 h(-1)). Total nitrogen uptake was also higher during the upwelling event (0.153-0.366 in June and 0.053-0.096 mu mol N L-1 h(-1) in September). Nitrogen uptake kinetics demonstrated a strong preference for ammonium and urea over nitrate in June.

Coccolithophore species as indicators of surface oceanographic conditions in the vicinity of Azores islands

During summer 2008 and spring 2009, surface oceanographic surveys were carried out around three islands of the Azores archipelago (Terceira, Sao Miguel and Santa Maria) to assess the phytoplankton distribution and associated physico-chemical processes. The Azores archipelago is a major feature in the biogeochemical North Atlantic Subtropical Gyre (NAST) province although its influence on the productivity of the surrounding ocean is poorly known. Surface phytoplankton was studied by microscopy and HPLC (High Precision Liquid Chromatography). The mean values for biomass proxy Chlorophyll a (Chla) ranged from 0.04 to 0.55 mu g L-1 (Chla maximum = 0.86 mu g L-1) and coccolithophores were the most abundant group, followed by small flagellates, Cyanobacteria, diatoms and dinoflagellates being the least abundant group. The distribution of phytoplankton and coccolithophore species in particular presented seasonal differences and was consistent with the nearshore influence of warm subtropical waters from the south Azores current and colder subpolar waters from the north. The satellite-derived circulation patterns showed southward cold water intrusions off Terceira and northward warm water intrusions off Santa Maria. The warmer waters signal was confirmed by the subtropical coccolithophore assemblage, being Discosphaera tubifera a constant presence under these conditions. The regions of enhanced biomass, either resulting from northern cooler waters or from island induced processes, were characterized by the presence of Emiliania huxleyi. Diatoms and dinoflagellates indicated coastal and regional processes of nutrient enrichment and areas of physical stability, respectively.

The response of phytoplankton to climate variability associated with the North Atlantic Oscillation

Inter-annual variability in the timing of phytoplankton spring bloom and phytoplankton community structure in the central North Atlantic Ocean was quantified using ocean color data and continuous plankton recorder (CPR) data. This variability was related to the North Atlantic Oscillation using correlation analysis and multivariate auto-regression models. The initiation of the spring bloom derived from CPR phytoplankton color index data is similar to that derived from satellite chlorophyll, and exhibits a nominal correlation with the sea surface temperature (SST) and the North Atlantic Oscillation (NAO). The extrapolated spring bloom timing suggested later initiation of blooms in the mid-1980s and earlier initiation of blooms in the 1990s. The climatological phytoplankton community structure in the central North Atlantic is dominated by diatoms, except for a shift in community composition favoring dinoflagellates in August. The ratio of diatoms to total phytoplankton abundance and the ratio of dinoflagellates to total phytoplankton abundance are both closely correlated with the NAO and SST. The extended time series of phytoplankton community structure between 1985 and 2009, deduced from the time series of SST and NAO over the same interval, showed a decadal shift away from diatoms towards dinoflagellates. The linkages between the NAO, and changes in stratification and phytoplankton processes occur over a larger scale than previously observed.