Véronique Rousseau

Calculating carbon biomass of Phaeocystis sp. from microscopic observations

Conversion factors for calculating carbon biomass ofPhaeocystis sp. colonies and free-living cells were determined from microscopic observations and chemical analysis conducted on cultured and naturalPhaeocystis sp. populations originating from the Southern Bight of the North Sea in 1986 and 1987. They allow calculation, in terms of carbon biomass, of the different forms ofPhaeocystis sp. that succeed each other when the population is growing, on the basis of microscopic observations. The latter include enumerations of free-living cells (flagellated and non-motile) and colonies, as well as colonial biovolume measurement. Specific application to natural populations from Dutch coastal waters during spring 1986 shows that more than 90% ofPhaeocystis sp. carbon biomass is under colonial form, most of it exceeding the grazing characteristics of current zooplankton at this period of the year. Detailed analysis of seasonal changes shows in addition that the size of the colonies greatly increases during the course ofPhaeocystis sp. flowering, reaching sizes as high as 1 mm diameter at the top of the bloom when nutrients are depleted. Physiologically this corresponds to an enhanced synthesis of mucilaginous substances, with the decrease of available nutrients leading to an increasing contribution of the matrix to the total colonial carbon during the course of the bloom. Carbon content ofPhaeocystis sp. colonies therefore greatly varies with their size, ranging from 0.3 to 1430 ngC colony−1.

The life cycle of Phaeocystis (Prymnesiophycaea): evidence and hypotheses

The present paper reviews the literature related to the life cycle of the prymnesiophyte Phaeocystis and its controlling factors and proposes novel hypotheses based on unpublished observations in culture and in the field. We chiefly refer to P. globosa Scherffel as most of the observations concern this species. P. globosa exhibits a complex alternation between several types of free-living cells (non-motile, flagellates, microzoopores and possibly macrozoospores) and colonies for which neither forms nor pathways have been completely identified and described. The different types of Phaeocystis cells were reappraised on the basis of existing microscopic descriptions complemented by unpublished flow cytometric investigations. This analysis revealed the existence of at least three different types of free-living cells identified on the basis of a combination of size, motility and ploidy characteristics: non-motile cells, flagellates and microzoospores. Their respective function within Phaeocystis life cycle, and in particular their involvement in colony formation is not completely understood. Observational evidence shows that Phaeocystis colonies are initiated at the early stage of their bloom each by one free-living cell. The mechanisms controlling this cellular transformation are still uncertain due to the lack of information on the overwintering Phaeocystis fomms and on the cell type responsible for colony induction. The existence of haploid microzoospores released from senescent colonies gives however some support to sexuality involvement at some stages of colony formation. Once colonies are formed, at least two mechanisms were identified as responsible of the spreading of colony form: colony multiplication by colonial division or budding and induction of new colony from colonial cells released in the external medium after colony disruption. The latter mechanism was clearly identified, involving at least two successive cell differentiations in the following sequence: motility development, subsequent flagella loss and settlement to a surface, mucus secretion and colony formation, colonial cell division and colony growth. Aggregate formation, cell motility development and subsequent emigration from the colonies, release of non-motile cells after colony lysis on the other hand, were identified as characteristics for termination of Phaeocystis colony development. These pathways were shown to occur similarly in natural environments. In the early stages of the bloom however, many recently-formed colonies were found on the setae of Chaetoceros spp, suggesting this diatom could play a key-role in Phaeocystis bloom inception. Analysis of the possible environmental factors regulating the transition between the different phases of the life cycle, suggested that nutrient status and requirement of a substrate for attachment of free-living cells would be essential for initiation of the colonial form. Physical constraints obviously would be important in determining colony shape and fragmentation although autogenic factors cannot be excluded. Some evidence exists that nutrients regulate colony division, while temperature and nutrient stress would stimulate cell emigration from the colonies.

Trophic efficiency of the planktonic food web in a coastal ecosystem dominated by Phaeocystis colonies

Abstract The trophic efficiency of the planktonic food web in the Phaeocystis-dominated ecosystem of the Belgian coastal waters was inferred from the analysis of the carbon flow network of the planktonic system subdivided into its different trophodynamic groups. A carbon budget was constructed on the basis of process-level field experiments conducted during the spring bloom period of 1998. Biomass and major metabolic activities of auto- and heterotrophic planktonic communities (primary production, bacterial production, nanoproto-, micro- and mesozooplankton feeding activities) were determined in nine field assemblages collected during spring at reference station 330. In 1998, the phytoplankton spring flowering was characterised by a moderate diatom bloom followed by a massive Phaeocystis colony bloom. Phaeocystis colonies, contributing 70% to the net primary production, escaped the linear food chain while the early spring diatom production supplied 74% of the mesozooplankton carbon uptake. The rest of mesozooplankton food requirement was, at the time of the Phaeocystis colony bloom, partially fulfilled by microzooplankton. Only one-third of the microzooplankton production, however, was controlled by mesozooplankton grazing pressure. Ungrazed Phaeocystis colonies were stimulating the establishment of a very active microbial network. On the one hand, the release of free-living cells from ungrazed colonies has been shown to stimulate the growth of microzooplankton, which was controlling 97% of the nanophytoplankton production. On the other hand, the disruption of ungrazed Phaeocystis colonies supplied the water column with large amounts of dissolved organic matter available for planktonic bacteria. The budget calculation suggests that ungrazed colonies contributed up to 60% to the bacterial carbon demand, while alternative sources (exudation, zooplankton egestion and lysis of other organisms) provided some 30% of bacterial carbon requirements. This suggests that the spring carbon demand of planktonic bacteria was satisfied largely by autogenic production. The trophic efficiency was defined as the ratio between mesozooplankton grazing on a given source and food production. In spite of its major contribution to mesozooplankton feeding, the trophic efficiency of the linear food chain, restricted to the grazing on diatoms, represented only 5.6% of the available net primary production. The trophic efficiency of the microbial food chain, the ratio between mesozooplankton grazing on microzooplankton and the resource inflow (the bacterial carbon demand plus the nanophytoplankton production) amounted to only 1.6%. These low trophic efficiencies together with the potential contribution of ungrazed Phaeocystis-derived production to the bacterial carbon demand suggest that during spring 1998 most of the Phaeocystis-derived production in the Belgian coastal area was remineralised in the water column.

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 vmax 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.

Coastal eutrophication of the Southern Bight of the North Sea : assessment and modelling

The most seriously threatened European seas are the North Sea, the Baltic and the Black Sea. Here the rapid degradation of these marine environments is examined comparatively to identify similarities and differences in the driving forces and responses of the marine systems. Although it is difficult to distinguish between anthropogenic changes and those due to natural climate fluctuations, an attempt is made here to define possible effects based on a multidisciplinary mix of recent observations and modelling. The systems studied range from almost totally enclosed domains to marginal seas adjoining a large ocean body, and from shallow, dissipative and tidally dominated shelf regions to deep, relatively stagnant basins with adjoining energetic shelf regions, and from completely anoxic to oxygen-saturated chemical environments. Climate control appears as a background theme, while the most significant human effect is eutrophication, which is leading to rapid changes in these systems.

Comparative analysis of food webs based on flow networks: effects of nutrient supply on structure and function of coastal plankton communities

The objective of COMWEB was to develop efficient analytical, numerical and experimental methods for assessing and predicting the effects of nutrient (N, P, Si) supply on the stability and persistence of pelagic food web structure and function in coastal waters. The experimental comparative work included a geographic gradient covering Baltic, Mediterranean, and NE Atlantic waters and a NE Atlantic gradient in state of eutrophication. COMWEB has been an experimental approach to coastal eutrophication, studying effects of enhanced nutrient supply on components and flows of the entire lower pelagic food web. Flow network representations of pelagic food webs has been a framework of data reduction and flows were established by sophisticated inverse modelling. Fundamental information on physiological properties of functional key species in the pelagic food web was used to constrain flow estimations. A main conclusion derived from the flow networks was that very little energy and materials were transferred from the microbial food web to the main food chain. The lower food web could therefore be described as two parallel food chains with relatively limited interaction between heterotrophic groups. Short-term effects of nutrient perturbations were examined in mesocosms along the geographic gradient. The response was comparable in all systems, with a stronger effect on the activity and biomass of autotrophic groups than those of heterotrophic ones. Mediterranean waters showed much lower autotrophic biomass response than Baltic and NE Atlantic waters, which responded almost equally. The response of primary production was, however, more comparable. High phytoplankton lysis rate explained this low accumulation of biomass in Mediterranean waters. The study of Atlantic coastal waters of different eutrophic states revealed that the ecological response was higher in the closed nutrient perturbed mesocosms than in open systems exposed for >4 summer months (summer/autumn season). The Atlantic lagoon evolved gradually from the natural oligotrophic situation towards the more eutrophicated North Sea during fertilisation. The responses observed on seasonal and long-term scale (>10 years) may therefore be equal. The differences between short-term (weeks) and intermediate-term (seasonal) responses is most likely a result of the different time scales of perturbation and observation and the variable exchange rates with surrounding waters (water dilution rate). The analysis of pelagic flow networks provided a framework of diagnostic criteria for state and quality assessment of coastal waters. The nutrient loading rates related better to estimates of biotic fluxes than to concentrations of biotic compartments and total nutrients. On the contrary, the concentration of biotic compartments, or the biomasses, related better to total nutrient concentrations. Primary production, mesozooplankton grazing and growth, fraction of primary production consumed by grazers, bacterial production relative to primary production, cycling indices, and path lengths were all well related to nutrient loading rate. Autotrophic biomass, ratio of autotrophic to heterotrophic biomass, and fraction of pico-cyanobacteria of total autotrophic biomass were all related to total nutrients. Some of these variables, which responded equally in all systems, have the potential of becoming unified response functions in a management model for European coastal waters. COMWEB has provided further insight into the mechanisms behind coastal eutrophication. A main achievement is the conceptual framework for unified response functions, important components of management models for nutrient emission to coastal waters.

Composition, assimilation and degradation of Phaeocystis globosa-derived fatty acids in the North Sea

The fate of a Phaeocystis globosa bloom in the southern North Sea off Belgium, the Netherlands and Germany in May 1995 was investigated during a cruise with RV ‘Belgica’. We used fatty acids as biomarkers to follow the fate of Phaeocystis-derived biomass of a Phaeocystis-dominated spring bloom. The bloom, in which up to >99% of the biomass was contributed by Phaeocystis, showed a fatty acid composition with a characteristically high abundance of polyunsaturated C18-fatty acids, which increased in concentration with number of double bonds up to 18:5 (n-3), and high concentrations of 20:5 (n-3) and 22:6 (n-3). In contrast to most previous studies, fatty acid analysis of the mesozooplankton community (mainly calanoid copepods) and meroplankton (Carcinus maenas megalope) indicated that P. globosa was a major component (ca. 70% and 50%, respectively) in the diet of these organisms. Massive accumulations of amorphous grey aggregates, in which Phaeocystis colonies were major components, were dominated by saturated fatty acids and contained only few of the polyunsaturated C18-fatty acids. A hydrophobic surface slick that covered the water surface during the bloom showed very similar patterns. Foam patches contained few Phaeocystis-typical fatty acids, but increased amounts of diatom-typical compounds such as 16:1 (n-7) and 20:5 (n-3), and 38% fatty alcohols, indicating that wax esters dominated the lipid fraction in the foam with ca. 76% (w/w). The fatty acid compositions of surface sediment showed that no sedimentation of fresh Phaeocystis occurred during the study. The results indicate that Phaeocystis-derived organic matter degraded while floating or in suspension, and had not reached the sediment in substantial amounts.

Optical properties of algal blooms in an eutrophicated coastal area and its relevance to remote sensing

The Southern Bight of the North Sea is characterised by a large influence of river inputs, which results in eutrophication of the area. High concentrations of plankton biomass and suspended matter have been reported for this area, in relation with blooms of different species and resuspension of bottom sediments. In spring the haptophyte Phaeocystis globosa blooms throughout the area reaching up to 30 mg Chlorophyll m-3 or more nearshore. This event is followed in June by red tides of the dinoflagellate Noctiluca scintillans. These blooms are concurrent with different species of diatoms. The strong optical signature of these blooms is clear to human observers making them potentially detectable in satellite imagery. As a first step in this direction, sampling has been carried out in the area, during Phaeocystis and Noctiluca blooms in 2003 and 2004. Phytoplankton pigments and inherent optical properties (particle, detrital and phytoplankton absorption) have been measured spectrophotometrically, and in situ using an ac-9 for total absorption and particle scattering. Field data were compared with optical properties of pure species obtained in laboratory. In parallel, water-leaving reflectance has been also measured. In this paper we characterise the optical signatures of diatoms, Phaeocystis and Noctiluca and their contribution to total absorption. The impact on water-leaving reflectance spectra is evaluated; in order to assess the conditions in which remote sensing can provide information for monitoring the timing, extent and magnitude of blooms in this coastal area.