Structure and diversity of intertidal benthic diatom assemblages in contrasting shores: a case study from the Tagus estuary1

The structure of intertidal benthic diatoms assemblages in the Tagus estuary was investigated during a 2-year survey, carried out in six stations with different sediment texture. Nonparametric multivariate analyses were used to characterize spatial and temporal patterns of the assemblages and to link them to the measured environmental variables. In addition, diversity and other features related to community physiognomy, such as size-class or life-form distributions, were used to describe the diatom assemblages. A total of 183 diatom taxa were identified during cell counts and their biovolume was determined. Differences between stations (analysis of similarity (ANOSIM), R=0.932) were more evident than temporal patterns (R=0.308) and mud content alone was the environmental variable most correlated to the biotic data (BEST, rho=0.863). Mudflat stations were typically colonized by low diversity diatom assemblages (H' similar to 1.9), mainly composed of medium-sized motile epipelic species (250-1,000 mu m(3)), that showed species-specific seasonal blooms (e.g., Navicula gregaria Donkin). Sandy stations had more complex and diverse diatom assemblages (H' similar to 3.2). They were mostly composed by a large set of minute epipsammic species (<250 mu m(3)) that, generally, did not show temporal patterns. The structure of intertidal diatom assemblages was largely defined by the interplay between epipelon and epipsammon, and its diversity was explained within the framework of the Intermediate Disturbance Hypothesis. However, the spatial distribution of epipelic and epipsammic life-forms showed that the definition of both functional groups should not be over-simplified.

Macroscale factors affecting diatom abundance: a synergistic use of Continuous Plankton Recorder and satellite remote sensing data

Diatoms exist in almost every aquatic regime; they are responsible for 20% of global carbon fixation and 25% of global primary production, and are regarded as a key food for copepods, which are subsequently consumed by larger predators such as fish and marine mammals. A decreasing abundance and a vulnerability to climatic change in the North Atlantic Ocean have been reported in the literature. In the present work, a data matrix composed of concurrent satellite remote sensing and Continuous Plankton Recorder (CPR) in situ measurements was collated for the same spatial and temporal coverage in the Northeast Atlantic. Artificial neural networks (ANNs) were applied to recognize and learn the complex non-monotonic and non-linear relationships between diatom abundance and spatiotemporal environmental factors. Because of their ability to mimic non-linear systems, ANNs proved far more effective in modelling the diatom distribution in the marine ecosystem. The results of this study reveal that diatoms have a regular seasonal cycle, with their abundance most strongly influenced by sea surface temperature (SST) and light intensity. The models indicate that extreme positive SSTs decrease diatom abundances regardless of other climatic conditions. These results provide information on the ecology of diatoms that may advance our understanding of the potential response of diatoms to climatic change.

Protection of cells from salinity stress by extracellular polymeric substances in diatom biofilms

Diatom biofilms are abundant in the marine environment. It is assumed (but untested) that extracellular polymeric substances(EPS), produced by diatoms, enable cells to cope with fluctuating salinity. To determine the protective role of EPS, Cylindrotheca closterium was grown in xanthan gum at salinities of 35, 50, 70 and 90 ppt. A xanthan matrix significantly increased cell viability (determined by SYTOX-Green), growth rate and population density by up to 300, 2, 300 and 200%, respectively. Diatoms grown in 0.75% w/v xanthan, subjected to acute salinity shock treatments (at salinities 17.5, 50, 70 and 90 ppt) maintained photosynthetic capacity, F_q′/F_m′, within 4% of pre-shock values, whereas F_q′/F_m′ in cells grown without xanthan declined by up to 64% with hypersaline shock. Biofilms that developed in xanthan at standard salinity helped cells to maintain function during salinity shock. These results provide evidence of the benefits of living in an EPS matrix for biofilm diatoms.

Interaction Effects of Light, Temperature and Nutrient Limitations (N, P and Si) on Growth, Stoichiometry and Photosynthetic Parameters of the Cold-Water Diatom Chaetoceros wighamii

Light (20-450 μmol photons m^-2 s^-1), temperature (3-11°C) and inorganic nutrient composition (nutrient replete and N, P and Si limitation) were manipulated to study their combined influence on growth, stoichiometry (C:N:P:Chl a) and primary production of the cold water diatom Chaetoceros wighamii. During exponential growth, the maximum growth rate (~0.8 d^-1) was observed at high temperture and light; at 3°C the growth rate was ~30% lower under similar light conditions. The interaction effect of light and temperature were clearly visible from growth and cellular stoichiometry. The average C:N:P molar ratio was 80:13:1 during exponential growth, but the range, due to different light acclimation, was widest at the lowest temperature, reaching very low C:P (~50) and N:P ratios (~8) at low light and temperature. The C:Chl a ratio had also a wider range at the lowest temperature during exponential growth, ranging 16-48 (weight ratio) at 3°C compared with 17-33 at 11°C. During exponential growth, there was no clear trend in the Chl a normalized, initial slope (α*) of the photosynthesis-irradiance (PE) curve, but the maximum photosynthetic production (P_m) was highest for cultures acclimated to the highest light and temperature. During the stationary growth phase, the stoichiometric relationship depended on the limiting nutrient, but with generally increasing C:N:P ratio. The average photosynthetic quotient (PQ) during exponential growth was 1.26 but decreased to <1 under nutrient and light limitation, probably due to photorespiration. The results clearly demonstrate that there are interaction effects between light, temperature and nutrient limitation, and the data suggests greater variability of key parameters at low temperature. Understanding these dynamics will be important for improving models of aquatic primary production and biogeochemical cycles in a warming climate.

Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea

We introduce a trait-based description of diatom functional diversity to an existing plankton functional type (PFT) model, implemented for the eutrophied coastal ecosystem in the Southern Bight of the North Sea. The trait-based description represents a continuum of diatom species, each characterized by a distinct cell volume, and includes size dependence of four diatom traits: the maximum growth rate, the half-saturation constants for nutrient uptake, the photosynthetic efficiency, and the relative affinity of copepods for diatoms. Through competition under seasonally varying forcing, the fitness of each diatom varies throughout time, and the outcome of competition results in a changing community structure. The predicted seasonal change in mean cell volume of the community is supported by field observations: smaller diatoms, which are more competitive in terms of resource acquisition, prevail during the first spring bloom, whereas the summer bloom is dominated by larger species which better resist grazing. The size-based model is used to determine the ecological niche of diatoms in the area and identifies a range of viable sizes that matches observations. The general trade-off between small, competitive diatoms and large, grazing-resistant species is a convenient framework to study patterns in diatom functional diversity. PFT models and trait-based approaches constitute promising complementary tools to study community structure in marine ecosystems.

Decrease in diatom palatability contributes to bloom formation in the Western English Channel

The aim of this paper is to investigate the role of phytoplankton nutritional status in the formation of the spring bloom regularly observed at the station L4 in the Western English Channel. Using a modelling approach, we tested the hypothesis that the increase in light from winter to spring induces a decrease in diatom nutritional status (i.e., an increase in the C:N and C:P ratios), thereby reducing their palatability and allowing them to bloom. To this end, a formulation describing the Stoichiometric Modulation of Predation (SMP) has been implemented in a simplified version of the European Regional Seas Ecosystem Model (ERSEM). The model was coupled with the General Ocean Turbulence Model (GOTM), implemented at the station L4 and run for 10 years (2000-.2009). Simulated carbon to nutrient ratios in diatoms were analysed in relation to microzooplankton biomass, grazing and assimilation efficiency. The model reproduced in situ data evolutions and showed the importance of microzooplankton grazing in controlling the early onset of the bloom. Simulation results supported our hypothesis and provided a conceptual model explaining the formation of the diatom spring bloom in the investigated area. However, additional data describing the microzooplankton grazing impact and the variation of carbon to nutrient ratios inside phytoplanktonic cells are required to further validate the proposed mechanisms.

Characterisation of algicidal bacterial exometabolites against the lipid-accumulating diatom Skeletonema sp.

Microalgae are of increasing interest due to their occurrence in the environment as harmful algal blooms and as a source of biomass for the production of fine and bulk chemicals. A method for the low cost disruption of algal biomass for environmental remediation or bioprocessing is desirable. Naturally-occurring algal lytic agents from bacteria could provide a cost-effective and environmentally desirable solution. A screen for algal lytic agents against a range of marine microalgae has identified two strains of algicidal bacteria isolated from the coastal region of the Western English Channel. Both strains (designated EC-1 and EC-2) showed significant algicidal activity against Skeletonema sp. and were identified as members of Alteromonas sp. and Maribacter sp. respectively. Characterisation of the two bioactivities revealed that they are small extracellular metabolites displaying thermal and acid stability. Purification of the EC-1 activity to homogeneity and initial structural analysis has identified it as a putative peptide with a mass of 1266. amu.

Characterisation of algicidal bacterial exometabolites against the lipid-accumulating diatom Skeletonema sp.

Microalgae are of increasing interest due to their occurrence in the environment as harmful algal blooms and as a source of biomass for the production of fine and bulk chemicals. A method for the low cost disruption of algal biomass for environmental remediation or bioprocessing is desirable. Naturally-occurring algal lytic agents from bacteria could provide a cost-effective and environmentally desirable solution. A screen for algal lytic agents against a range of marine microalgae has identified two strains of algicidal bacteria isolated from the coastal region of the Western English Channel. Both strains (designated EC-1 and EC-2) showed significant algicidal activity against Skeletonema sp. and were identified as members of Alteromonas sp. and Maribacter sp. respectively. Characterisation of the two bioactivities revealed that they are small extracellular metabolites displaying thermal and acid stability. Purification of the EC-1 activity to homogeneity and initial structural analysis has identified it as a putative peptide with a mass of 1266. amu.

A role for diatom-like silicon transporters in calcifying coccolithophores.

Biomineralization by marine phytoplankton, such as the silicifying diatoms and calcifying coccolithophores, plays an important role in carbon and nutrient cycling in the oceans. Silicification and calcification are distinct cellular processes with no known common mechanisms. It is thought that coccolithophores are able to outcompete diatoms in Si-depleted waters, which can contribute to the formation of coccolithophore blooms. Here we show that an expanded family of diatom-like silicon transporters (SITs) are present in both silicifying and calcifying haptophyte phytoplankton, including some globally important coccolithophores. Si is required for calcification in these coccolithophores, indicating that Si uptake contributes to the very different forms of biomineralization in diatoms and coccolithophores. Significantly, SITs and the requirement for Si are absent from highly abundant bloom-forming coccolithophores, such as Emiliania huxleyi. These very different requirements for Si in coccolithophores are likely to have major influence on their competitive interactions with diatoms and other siliceous phytoplankton.

A role for diatom-like silicon transporters in calcifying coccolithophores.

Biomineralization by marine phytoplankton, such as the silicifying diatoms and calcifying coccolithophores, plays an important role in carbon and nutrient cycling in the oceans. Silicification and calcification are distinct cellular processes with no known common mechanisms. It is thought that coccolithophores are able to outcompete diatoms in Si-depleted waters, which can contribute to the formation of coccolithophore blooms. Here we show that an expanded family of diatom-like silicon transporters (SITs) are present in both silicifying and calcifying haptophyte phytoplankton, including some globally important coccolithophores. Si is required for calcification in these coccolithophores, indicating that Si uptake contributes to the very different forms of biomineralization in diatoms and coccolithophores. Significantly, SITs and the requirement for Si are absent from highly abundant bloom-forming coccolithophores, such as Emiliania huxleyi. These very different requirements for Si in coccolithophores are likely to have major influence on their competitive interactions with diatoms and other siliceous phytoplankton.

Unexpected prevalence of parasite 18S rDNA sequences in winter among Antarctic marine protists

Parasites are not typically considered to be important components of polar marine ecosystems. It was therefore surprising when 18S rDNA surveys of protists in the West Antarctic Peninsula in winter revealed high abundances of parasite sequences. Parasite sequences made up, on average, over half (52%) of sequence reads in samples from deep water in winter. Winter surface water and sediment samples contained relatively fewer, but still strikingly high, parasite sequence reads (13 and 9%, respectively), while surface water samples in summer contained fewer parasite sequences (1.8%). A total of 1028 distinct parasite Operational Taxonomic Units were observed in winter, with the largest abundances and diversities within Syndiniales groups I and II, including Amoebophrya. Less abundant parasite sequence groups included Apicomplexa, Blastodinium, Chytriodinium, Cryptocaryon, Paradinium, Perkinsidae, Pirsonia and Ichthyophonae. Parasite sequence distributions suggested interactions with known hosts, such as diatom parasites which were mainly in the sediments, where resting spores of Chaetoceros spp. diatoms were abundant. Syndiniales sequences were correlated with radiolarian sequences, suggesting parasite–host interactions. The abundant proportions of parasite sequences indicate a potentially important role for parasites in the Antarctic marine ecosystem, with implications for plankton population dynamics, the role of the microbial loop, carbon flows and ecosystem responses to ongoing anthropogenic climate change.