A mathematical model was developed to represent the dynamics of predation and assimilation of ingested material by heterotrophic marine micro-zooplankton. Predation rate was made a rectangular hyperbolic function of prey carbon (C)... more
A mathematical model was developed to represent the dynamics of predation and assimilation of ingested material by heterotrophic marine micro-zooplankton. Predation rate was made a rectangular hyperbolic function of prey carbon (C) concentration modified to simulate the prey selectivity that these organisms have been observed experimentally to exhibit in response to prey nutritional quality. We chose prey N:C ratio as an index of selectivity and related both the maximum predation rate (PM) and the assimilation efficiency (AE) to changes in prey nitrogen:carbon (N:C). Changes in PM simulated the phenomena of "surge feeding" and "prey rejection". Changes in AE simulated sub-optimal assimilation of material from ingested prey, which in turn, served to decrease the ingestion rate on this material. The model was parameterised using laboratory data sets that followed ingestion of non-growing phytoplankton prey of different N:C ratio by a microflagellate predator. Investigative simulations in transient conditions, incorporating growing prey and hence with changing N:C ratio, indicated both PM and AE influenced the quantitative and temporal dynamics of C transfer to the higher trophic level. In particular, we noted that although various model formulations predicted similar trophic transfer of C, this was achieved on very different time scales. The inclusion of heterotrophic micro-zooplankton within food web models is a necessary step in accurate prediction of pelagic nutrient flux but requires physiological models carefully parameterised and tested in both steady state and transient conditions to ensure accurate simulation.
There is a perception that anthropogenically-driven changes in nutrient supply to coastal waters influences the abundance, frequency and toxicity of harmful algal blooms (HABs) through a change in the form or ratio of nutrient that limits... more
There is a perception that anthropogenically-driven changes in nutrient supply to coastal waters influences the abundance, frequency and toxicity of harmful algal blooms (HABs) through a change in the form or ratio of nutrient that limits phytoplankton growth. If nutrient concentrations are not limiting for growth, then ratios do not influence floristic composition. At non-limiting concentrations, evidence that alteration of nitrogen: phosphorus (N:P) ratios has stimulated HABs is limited, and primarily based on hypothesised relationships in relatively few locations (in particular: Tolo Harbour Hong Kong and Dutch Coastal Waters). In all cases, an unequivocal causal link between an increase in HABs (frequency, magnitude or duration) and change in N or P as the limiting nutrient is difficult to establish. The silicon (Si) limitation hypothesis is generally supported by experimental evidence and field data on the nuisance flagellate Phaeocystis. We found little evidence that high N:Si ratios preferentially promote harmful dinoflagellates over benign species. Laboratory studies demonstrate that nutrient ratios can influence toxin production, but genus and species specific differences and environmental control make extrapolation of these data to the field difficult. Studies of the role of dissolved and particulate organic nutrients in the growth of HAB species, while limited, demonstrate the potential for organic nutrients (especially organic N) to support the growth of a range of HAB species. There is a clear need for better understanding of the role of mixotrophy in the formation of HABs and for studies of HAB and non-HAB species in competition for environmentally realistic concentrations of organic nutrients.
Variable inorganic nutrient concentrations were used to generate silicate (Si) or nitrogen (N) limited conditions in cultures of the marine diatom Skeletonema costatum. Dissolved organic matter (DOM) harvested in the nutrient limited... more
Variable inorganic nutrient concentrations were used to generate silicate (Si) or nitrogen (N) limited conditions in cultures of the marine diatom Skeletonema costatum. Dissolved organic matter (DOM) harvested in the nutrient limited phase of these cultures was added to a natural bacterial community. Enhanced bacterial abundance and bacterial production were observed, in comparison to un-supplemented controls, when Si-limited (Si-DOM) rather than N-limited (N-DOM) diatom derived DOM or inorganic nutrients was added. This indicates that the bacterial population was limited by organic rather than inorganic resources but only Si-DOM had sufficient lability to alleviate this. Within the bacterial assemblage, a notable increase in the proportion of γ-Proteobacteria was evident on receipt of only Si-DOM. Assessment of the composition of the added DOM suggested that the observed dynamics were related to organic matter composition rather than molecular size as, within the Si-and N-DOM, the proportions of low and high molecular weight compounds were similar, but the polysaccharide and protein signatures were different.
Protists are major consumers of phytoplankton and bacterioplankton in the marine environment, playing a key role in carbon cycling and nutrient regeneration. Consequently, the feeding activities of planktonic protists have been... more
Protists are major consumers of phytoplankton and bacterioplankton in the marine environment, playing a key role in carbon cycling and nutrient regeneration. Consequently, the feeding activities of planktonic protists have been intensively studied. In particular, the dinoflagellate, Oxyrrhis marina, has been used extensively as a model predator in laboratory-based feeding studies. This is predominately because it is easy to culture and manipulate and also because its plasticity in feeding behaviour allows it to represent a broad range of free-living heterotrophic protists. Prey preferences of this versatile, but selective, predator have been well documented. However, our ability to determine the factors driving prey selectivity is limited by a poor understanding of the detailed mechanisms underlying O. marina feeding. Within this study, we explore the known aspects of O. marina feeding behaviour and highlight areas that require further research to exploit this species as an effective model protist predator.
Many "model" protists are maintained in culture and used, experimentally, to answer questions associated with planktonic processes. Given the current interest and rapidly increasing amount of literature on the heterotrophic dinoflagellate... more
Many "model" protists are maintained in culture and used, experimentally, to answer questions associated with planktonic processes. Given the current interest and rapidly increasing amount of literature on the heterotrophic dinoflagellate Oxyrrhis marina, we present in this special issue a series of focused, interlinked research articles. Being written by experts in their respective fields, the authors have included unpublished data and in all cases have offered a synthesis of data and ideas. Furthermore, we have encouraged cross-paper discourse, emphasizing the interdisciplinary nature of our work and the utility of O. marina to this end; we also offer guidance, both practical and intellectual, on how future research related to O. marina might progress. In this introduction, however, we raise the wider issue of which criteria are required to consider a taxon as a "model species". We then assess the extent to which O. marina can fill this role. In general, we recognize O. marina as a model in three distinct disciplines: ecology, evolution/genomics and biogeography. Of possibly greater importance, we recognize that if O. marina continues to be studied at an escalating rate, there will be a concomitant increase in realized and potential synergies across these fields.
A nutrient-manipulation mesocosm study was conducted in Norwegian coastal waters to determine the effect on a nitrogen-limited microbial food web of changes in the relative concentrations of dissolved organic and inorganic nitrogen (N).... more
A nutrient-manipulation mesocosm study was conducted in Norwegian coastal waters to determine the effect on a nitrogen-limited microbial food web of changes in the relative concentrations of dissolved organic and inorganic nitrogen (N). Four replicated treatments were studied: no N addition, inorganic N, organic N, or a 50 : 50 mix of inorganic and organic N. Comprehensive abundance, biomass, and rate measurements were made over the 20-d experiment. The form of N available influenced species composition, succession, and the efficiency of carbon (C) incorporation. Inorganic N generated a bloom of the diatom Leptocylindrus danicus, even though silicon concentrations were below 1 mmol L 21 . The c-proteobacteria initially competed effectively with diatoms for inorganic N, exhibiting high productivity and escaping grazing control. Organic N, when available alone, was utilized more slowly by the microbial community, with diatom growth, fueled by regenerated N, achieving a lower peak abundance. A more diverse diatom species assemblage was also evident. The c-proteobacteria were less prevalent and net bacterial productivity was initially lower when the N source was organic and, in contrast to the response following inorganic N addition, the bacterial community was initially grazer controlled. A rapid succession of heterotrophic grazers occurred but with differences in species and their contribution to biomass between treatments. The efficiency of C-biomass production was lower when the N source was purely organic. However, when both inorganic and organic N were available, biomass yield was greater than the mean of the two single N source treatments. 1 Corresponding author ([email protected]).
The time course of colored dissolved organic matter (CDOM) absorption and fluorescence were monitored during 50 to 70 days of laboratory incubations with water collected in the coastal upwelling system of the Ría de Vigo (NW Iberian... more
The time course of colored dissolved organic matter (CDOM) absorption and fluorescence were monitored during 50 to 70 days of laboratory incubations with water collected in the coastal upwelling system of the Ría de Vigo (NW Iberian Peninsula) under contrasting hydrographic conditions. CDOM fluorescence at peak-T (Ex/Em, 280/350 nm), characteristic of protein-like materials, decayed at a 1st order degradation rate constant (k T ) of 0.28± 0.13 day − 1 (average ± SD). k T covaried (R 2 = 0.86, p b 0.0002) with the degradation rate of the bulk DOC (k DOC ), but the protein-like materials degraded 72 ± 23% faster than DOC. Therefore, this study confirms that the CDOM fluorescence at peak-T can be used as a proxy to a DOM fraction significantly more labile than the bulk bioavailable DOC. In parallel with the decay of DOC and protein-like fluorescence, an increase in CDOM fluorescence at peak-M (Ex/Em, 320/410 nm) during the course of the incubations verified the production of marine humic-like substances as a by-product of the microbial metabolism. CDOM fluorescence at peak-M built-up at a production rate (k M ) of 0.06 ± 0.01 day − 1 (average ± SD) in the Ría de Vigo. Furthermore, the slope of the linear regression between k DOC and k M (R 2 = 0.64, p b 0.001) revealed that the formation of marine humic-like substances occurred at about one fifth of the rate of net DOC consumption.
Phytoplankton underpin the marine food web in shelf seas, with some species having properties that are harmful to human health and coastal aquaculture. Pressures such as climate change and anthropogenic nutrient input are hypothesized to... more
Phytoplankton underpin the marine food web in shelf seas, with some species having properties that are harmful to human health and coastal aquaculture. Pressures such as climate change and anthropogenic nutrient input are hypothesized to influence phytoplankton community composition and distribution. Yet the primary environmental drivers in shelf seas are poorly understood. To begin to address this in North Western European waters, the phytoplankton community composition was assessed in light of measured physical and chemical drivers during the ''Ellett Line'' cruise of autumn 2001 across the Scottish Continental shelf and into adjacent open Atlantic waters. Spatial variability existed in both phytoplankton and environmental conditions, with clear differences not only between on and off shelf stations but also between different on shelf locations. Temperature/salinity plots demonstrated different water masses existed in the region. In turn, principal component analysis (PCA), of the measured environmental conditions (temperature, salinity, water density and inorganic nutrient concentrations) clearly discriminated between shelf and oceanic stations on the basis of DIN:DSi ratio that was correlated with both salinity and temperature. Discrimination between shelf stations was also related to this ratio, but also the concentration of DIN and DSi. The phytoplankton community was diatom dominated, with multidimensional scaling (MDS) demonstrating spatial variability in its composition. Redundancy analysis (RDA) was used to investigate the link between environment and the phytoplankton community. This demonstrated a significant relationship between community composition and water mass as indexed by salinity (whole community), and both salinity and DIN:DSi (diatoms alone). Diatoms of the Pseudo-nitzschia seriata group occurred at densities potentially harmful to shellfish aquaculture, with the potential for toxicity being elevated by the likelihood of DSi limitation of growth at most stations and depths.
The recently completed European Census of Marine Life, conducted within the framework of the global Census of Marine Life programme (2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008)(2009)(2010), markedly enhanced our understanding of... more
The recently completed European Census of Marine Life, conducted within the framework of the global Census of Marine Life programme (2000)(2001)(2002)(2003)(2004)(2005)(2006)(2007)(2008)(2009)(2010), markedly enhanced our understanding of marine biodiversity in European Seas, its importance within ecological systems, and the implications for human use. Here we undertake a synthesis of present knowledge of biodiversity in European Seas and identify remaining challenges that prevent sustainable management of marine biodiversity in one of the most exploited continents of the globe. Our analysis demonstrates that changes in faunal standing stock with depth depends on the size of the fauna, with macrofaunal abundance only declining with increasing water depth below 1000 m, whilst there was no obvious decrease in meiofauna with increasing depth. Species richness was highly variable for both deep water macro-and meio-fauna along latitudinal and longitudinal gradients. Nematode biodiversity decreased from the Atlantic into the Mediterranean whilst latitudinal related biodiversity patterns were similar for both faunal groups investigated, suggesting that the same environmental drivers were influencing the fauna. While climate change and habitat degradation are the most frequently implicated stressors affecting biodiversity throughout European Seas, quantitative understanding, both at individual and cumulative/synergistic level, of their influences are often lacking. Full identification and quantification of species, in even a single marine habitat, remains a distant goal, as we lack integrated data-sets to quantify these. While the importance of safeguarding marine biodiversity is recognised by policy makers, the lack of advanced understanding of species diversity and of a full survey of any single habitat raises huge challenges in quantifying change, and facilitating/prioritising habitat/ecosystem protection. Our study highlights a pressing requirement for more complete biodiversity surveys to be undertaken within contrasting habitats, together with investigations in biodiversity-ecosystem functioning links and identification of separate and synergistic/cumulative humaninduced impacts on biodiversity.
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Laboratory experiments were conducted to study nitrogen (N) regeneration by the heterotrophic marine dinoflagellate Oxyrrhis marina when ingesting phytoplankton prey of two different species and of two alternative carbon:nitrogen (C:N)... more
Laboratory experiments were conducted to study nitrogen (N) regeneration by the heterotrophic marine dinoflagellate Oxyrrhis marina when ingesting phytoplankton prey of two different species and of two alternative carbon:nitrogen (C:N) ratios. Experiments were conducted in the presence of L-methionine sulfoximine (MSX) which acts as a glutamine synthetase inhibitor. Utilisation by phytoplankton of N regenerated by protozoans and other organisms drives secondary production in marine food webs. However, the rapid utilisation of this N by phytoplankton has previously hampered accurate assessment of the efficiency of protozoan N regeneration. This phenomenon is particularly problematic when the phytoplankton are nutrient stressed and most likely to rapidly utilise N. The use of MSX prevented significant utilisation by phytoplankton of protozoan regenerated N. Hence, by removing the normal pathway of N cycling, we were able to determine the N regeneration efficiency (NRE) of the protozoan. The results suggested that predator NRE could be explained in terms of the relative CN stoichiometry of prey and predator. Using a mathematical model we demonstrated that changing the method used to simulate the NRE of the protozoan trophic level has the potential to markedly modify the predicted dynamics of the simulated microbial food web.
An investigation into the diversity of the dinoflagellate Alexandrium was carried out during August 2007 within two fjordic sea lochs in the Shetland Isles, Scotland. The co-occurrence in the water column of the non-toxic West European... more
An investigation into the diversity of the dinoflagellate Alexandrium was carried out during August 2007 within two fjordic sea lochs in the Shetland Isles, Scotland. The co-occurrence in the water column of the non-toxic West European (W.E. or Gr.III) and the neurotoxic North American (N.A. or Gr.I) ribotypes of A. tamarense was demonstrated using fluorescent in situ hybridisation. A patch of A. tamarense (W.E.) localised at $10 m depth and extending over 6 km was detected in 'Clift Sound' with concentrations locally reaching $1 Â 10 4 cells l À1 . A. tamarense (N.A.) was also observed there but despite the presence of toxins in net haul samples collected locally, concentrations were low and near limits of detection. Alexandrium concentrations were $1.5 Â 10 3 cells l À1 in 'Vaila Sound', where both W.E. and N.A. ribotypes were detected with equal relative abundances in some samples. Given the patchiness of A. tamarense populations and their possible organisation in thin layer structures, better vertical resolution through fine-scale sampling will be necessary for population dynamic studies. Implications for the shellfish industry are substantial since harmful microalgae patches may not be detected during routine monitoring. Moreover, the co-occurrence of morphologically indistinct toxic and non-toxic ribotypes will necessitate implementing molecular methods for their discrimination.
- by Keith Davidson and +2
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- Biodiversity
High-biomass harmful algal blooms can kill farmed fish through toxicity, physical effects or de-oxygenation of the water column. These blooms often form over spatially large areas meaning that Earth observation is well placed to monitor... more
High-biomass harmful algal blooms can kill farmed fish through toxicity, physical effects or de-oxygenation of the water column. These blooms often form over spatially large areas meaning that Earth observation is well placed to monitor and study them. In this letter, we present a statistical-based background subtraction technique that has been modified to detect high-biomass algal blooms. The method builds upon previous work and uses a statistical framework to combine spatial and temporal information to produce maps of bloom extent. Its statistical nature allows the approach to characterize the region of interest meaning that region-specific tuning is not needed. The accuracy of the approach has been evaluated using Moderate Resolution Imaging Spectroradiometer (MODIS) data and an in situ cell concentration dataset, resulting in a correct classification rate of 68.0% with a false alarm rate of 0.24 (n = 25). The method is then used to study the surface coverage of a large high-biomass harmful algal bloom of Karenia mikimotoi. The approach shows promise for the early warning of spatially large high-biomass algal blooms, providing valuable information to support in situ sampling campaigns.
Harmful algal blooms (HABs) are thought to be increasing in coastal waters worldwide. Anthropogenic nutrient enrichment has been proposed as a principal causative factor of this increase through elevated inorganic and/or organic nutrient... more
Harmful algal blooms (HABs) are thought to be increasing in coastal waters worldwide. Anthropogenic nutrient enrichment has been proposed as a principal causative factor of this increase through elevated inorganic and/or organic nutrient concentrations and modified nutrient ratios. We assess: 1) the level of understanding of the link between the amount, form and ratio of anthropogenic nutrients and HABs; 2) the evidence for a link between anthropogenically generated HABs and negative impacts on human health; and 3) the economic implications of anthropogenic nutrient/HAB interactions. We demonstrate that an anthropogenic nutrient-HAB link is far from universal, and where it has been demonstrated, it is most frequently associated with high biomass rather than low biomass (biotoxin producing) HABs. While organic nutrients have been shown to support the growth of a range of HAB species, insufficient evidence exists to clearly establish if these nutrients specifically promote the growth ...
High biomass Harmful Algal Blooms (HABs) such as Karenia mikimotoi and shellfish toxin producing HAB species continue to be observed in UK and Republic of Ireland waters. Regional differences continue to be seen in the distribution of... more
High biomass Harmful Algal Blooms (HABs) such as Karenia mikimotoi and shellfish toxin producing HAB species continue to be observed in UK and Republic of Ireland waters. Regional differences continue to be seen in the distribution of HABs in UK and RoI waters with impacts mainly observed in the south and west coast of Ireland and regions in the UK with a strong Atlantic influence e.g. Regions 1, 3, 4, 6 and 7. There is little monitoring aside from the continuous plankton recorder (CPR) in Region 8. The impacts from HABs in Wales, Northern Ireland and the Isle of Man are generally low.
- by Keith Davidson and +3
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Many "model" protists are maintained in culture and used, experimentally, to answer questions associated with planktonic processes. Given the current interest and rapidly increasing amount of literature on the heterotrophic dinoflagellate... more
Many "model" protists are maintained in culture and used, experimentally, to answer questions associated with planktonic processes. Given the current interest and rapidly increasing amount of literature on the heterotrophic dinoflagellate Oxyrrhis marina, we present in this special issue a series of focused, interlinked research articles. Being written by experts in their respective fields, the authors have included unpublished data and in all cases have offered a synthesis of data and ideas. Furthermore, we have encouraged cross-paper discourse, emphasizing the interdisciplinary nature of our work and the utility of O. marina to this end; we also offer guidance, both practical and intellectual, on how future research related to O. marina might progress. In this introduction, however, we raise the wider issue of which criteria are required to consider a taxon as a "model species". We then assess the extent to which O. marina can fill this role. In general, we recognize O. marina as a model in three distinct disciplines: ecology, evolution/genomics and biogeography. Of possibly greater importance, we recognize that if O. marina continues to be studied at an escalating rate, there will be a concomitant increase in realized and potential synergies across these fields.
The co-occurrence of morphologically identical toxic and non-toxic ribotypes of the biotoxin producing marine dinoflagellate Alexandrium tamarense presents a significant problem for its identification and enumeration, particularly in a... more
The co-occurrence of morphologically identical toxic and non-toxic ribotypes of the biotoxin producing marine dinoflagellate Alexandrium tamarense presents a significant problem for its identification and enumeration, particularly in a regulatory monitoring context. To address this, we have developed a fluorescence in situ hybridization-flow cytometry (FISH-FC) based method of cell identification and enumeration. This employed the taxa specific oligonucleotide probes TamToxC and TamA to fluorescently label (with the fluorochromes CY.3 and FITC) Group I (toxic) and Group III (non-toxic) A. tamarense ribotypes respectively. Detection was by fluorescence activated flow cytometric analysis. The FISH-FC method allowed effective discrimination between laboratory cultures of Group I and Group III ribotypes, with toxic and non-toxic cells creating distinct, easily identifiable, clusters in a flow cytometer bi-plot of side scatter (SSC) versus the green (FL1) fluorescence detection channel. ...