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The association of Phaeocystis spp. with small pennate diatoms during three Phaeocystis -dominated spring blooms were investigated in the Eastern English Channel (2003 and 2004) and in coastal waters of Western Norway during a mesocosm experiment (2005). In each of these studies, colonization of the surface of large Phaeocystis spp. colonies by small needle-shaped diatoms ( Pseudo-nitzschia spp.) were observed. In the English Channel the diatom Pseudo-nitzschia delicatissima colonized the surface of large (>100 μm) Phaeocystis globosa colonies. The abundance of Pseudo-nitzschia delicatissima reached 130 cells per colony and formed up to 70% of the total carbon associated with Phaeocystis cells during late bloom stages. In Norwegian waters, the surface of large (>250 μm) Phaeocystis pouchetii colonies were colonized by Pseudo-nitzschia cf. granii var. curvata and to a lesser degree by other phytoplankton and protist species, although the abundance of these diatoms was never greater than 40 cells per colony. Based on these observations we suggest that diatoms utilize Phaeocystis colonies not only as habitat, but that they are able to utilize the colonial matrix as a growth substrate. Furthermore, these observations indicate that a considerable fraction of biomass (chlorophyll) associated with Phaeocystis colonies, especially large colonies concerned with intense and prolonged blooms, are due to co-occurring plankton species and not exclusively Phaeocystis cells.

The worldwide colony-forming haptophyte phytoplankton Phaeocystis spp. are key organisms in trophic and biogeochemical processes in the ocean. Many organisms from protists to fish ingest cells and/or colonies of Phaeocystis . Reports on specific mortality of Phaeocystis in natural plankton or mixed prey due to grazing by zooplankton, especially protozooplankton, are still limited. Reported feeding rates vary widely for both crustaceans and protists feeding on even the same Phaeocystis types and sizes. Quantitative analysis of available data showed that: (1) laboratory-derived crustacean grazing rates on monocultures of Phaeocystis may have been overestimated compared to feeding in natural plankton communities, and should be treated with caution; (2) formation of colonies by P. globosa appeared to reduce predation by small copepods (e.g., Acartia , Pseudocalanus , Temora and Centropages ), whereas large copepods (e.g., Calanus spp.) were able to feed on colonies of Phaeocystis pouchetii ; (3) physiological differences between different growth states, species, strains, cell types, and laboratory culture versus natural assemblages may explain most of the variations in reported feeding rates; (4) chemical signaling between predator and prey may be a major factor controlling grazing on Phaeocystis ; (5) it is unclear to what extent different zooplankton, especially protozooplankton, feed on the different life forms of Phaeocystis in situ. To better understand the mechanisms controlling zooplankton grazing in situ, future studies should aim at quantifying specific feeding rates on different Phaeocystis species, strains, cell types, prey sizes and growth states, and account for chemical signaling between the predator and prey. Recently developed molecular tools are promising approaches to achieve this goal in the future.

A two-dimensional microscale (5 cm resolution) sampler was used over the course of a phytoplankton spring bloom dominated by Phaeocystis globosa to investigate the structural properties of chlorophyll a and seawater excess viscosity distributions. The microscale distribution patterns of chlorophyll a and excess viscosity were never uniform nor random. Instead they exhibited different types and levels of aggregated spatial patterns that were related to the dynamics of the bloom. The chlorophyll a and seawater viscosity correlation patterns were also controlled by the dynamics of the bloom with positive and negative correlations before and after the formation of foam in the turbulent surf zone. The ecological relevance and implications of the observed patchiness and biologically induced increase in seawater viscosity are discussed and the combination of the enlarged colonial form and mucus secretion is suggested as a competitive advantage of P. globosa in highly turbulent environments where this species flourishes.

Chemical defence is a potential mechanism contributing to the success of Phaeocystis species that repeatedly dominate the phytoplankton in coastal areas. Species within the genus Phaeocystis have long been suspected of imposing negative effects on co-occurring organisms. Recently a number of toxins have been extracted and identified from Phaeocystis samples, but it is not clear if they do enhance the competitive advantage of Phaeocystis species. In the present study the cytotoxic impact of live Phaeocystis pouchetii to human blood cells in close proximity, regardless of the nature of the responsible mechanism, was quantified using a bioassay. Haemolytic activity was measured during blooms of P. pouchetii in mesocosms. These environments were chosen to mimic natural conditions including chemically mediated interactions that could trigger defensive and/or allelopathic responses of Phaeocystis . Haemolytic activity correlated with P. pouchetii numbers and was absent during the preceding diatom bloom. Samples containing live P. pouchetii cells showed the highest activity, while filtered sea water and cell extracts were less haemolytic or without effect. Dose-response curves were linear up to 70% lysis, and haemolysis in samples containing live P. pouchetii cells reached EC_50 values comparable to known toxic prymnesiophytes (1.9 * 10^7 cells l^−1). Haemolytic activity was enhanced by increased temperature and light. The results indicate that unprotected and thus presumably vulnerable cells present in a P. pouchetii bloom may lyse within days.

Over the years, viruses have been shown to be mortality agents for a wide range of phytoplankton species, including species within the genus Phaeocystis (Prymnesiophyceae). With its polymorphic life cycle, its worldwide distribution, and the capacity of several of the Phaeocystis species to form dense blooms, this genus is a key player for our understanding of biogeochemical cycling of elements. This paper provides an overview of what is know to date about the ecological role of viruses in regulating Phaeocystis population dynamics. It explores which variables affect the algal host–virus interactions, and examines the impact of virally induced cell lysis of Phaeocystis on the function and structure of the pelagic food web as well as on the flow of organic carbon and nutrients.

Phaeocystis spp. cell and colony mass fluxes and their contribution to the vertical particulate organic carbon (POC) export from a wide range of stations were quantified by short-term sediment traps. The compilation of available data, ranging from polar to sub-arctic and boreal regions, revealed that Phaeocystis colonial and single cells frequently are observed in shallow sediment traps at 30–50 m depth (average of 7 ± 11% of POC export). A strong vertical export decline between 40 m and 100 m diminished the contribution of Phaeocystis spp. cell carbon to vertical export of POC to only 3 ± 2% at 100 m depth, with two exceptions (deeper mixed stations). Estimates of potential corresponding mucus contribution increased the average Phaeocystis spp. contribution to <5% of POC export. The vertical flux attenuation efficiency is higher for Phaeocystis spp. than for diatoms. The overall contribution of Phaeocystis spp. to vertical carbon export based on direct investigations of vertical organic carbon export is small.

We report data of a naturally occurring radionuclide, ^234Th, an in situ tracer, to investigate vertical export of biogenic matter during a vernal bloom of Phaeocystis pouchetii in the fjords of northern Norway. To optimise sampling of different stages of the bloom, three fjords with increasing oceanic influence (Balsfjord, Malangen fjord and Ullsfjord, respectively) were investigated in April 1997. Contrasting situations were encountered between the three fjords: the proliferation of P. pouchetii in Ullsfjord surface waters coincided with a drastic reduction of particulate ^234Th fluxes in traps, although particulate organic carbon (POC) and dimethylsulphoniopropionate (DMSP) were exported and ^234Th was available in surface waters. When large colonies make up a significant fraction of the vertical flux, as observed in Ullsfjord in April 1997, there may be a large and rapid change in the POC/^234Th ratio, further complicating the use of ^234Th as a tracer for POC export. The results suggest that the proliferation of Phaeocystis pouchetii during vernal bloom could temporary increase OC/^234Th ratio of particles and delay the particulate export of ^234Th, and probably of other particle-reactive species, from surface waters.

Seawater concentrations of the climate-cooling, volatile sulphur compound dimethylsulphide (DMS) are the result of numerous production and consumption processes within the marine ecosystem. Due to this complex nature, it is difficult to predict temporal and geographical distribution patterns of DMS concentrations and the inclusion of DMS into global ocean climate models has only been attempted recently. Comparisons between individual model predictions, and ground-truthing exercises revealed that information on the functional relationships between physical and chemical ecosystem parameters, biological productivity and the production and consumption of DMS and its precursor dimethylsulphoniopropionate (DMSP) is necessary to further refine future climate models. In this review an attempt is made to quantify these functional relationships. The description of processes includes: (1) parameters controlling DMSP production such as species composition and abiotic factors; (2) the conversion of DMSP to DMS by algal and bacterial enzymes; (3) the fate of DMSP-sulphur due to, e.g., grazing, microbial consumption and sedimentation and (4) factors controlling DMS removal from the water column such as microbial consumption, photo-oxidation and emission to the atmosphere. We recommend the differentiation of six phytoplankton groups for inclusion in future models: eukaryotic and prokaryotic picoplankton, diatoms, dinoflagellates, and other phytoflagellates with and without DMSP-lyase activity. These functional groups are characterised by their cell size, DMSP content, DMSP-lyase activity and interactions with herbivorous grazers. In this review, emphasis is given to ecosystems dominated by the globally relevant haptophytes Emiliania huxleyi and Phaeocystis sp., which are important DMS and DMSP producers.

Dimethyl sulphide (DMS) is a biogenic gas of climatic significance on which limited information is available from the Indian Ocean. To fill this gap, we collected data on DMS and total dimethylsulphoniopropionate (DMSP_t) by participating in a dozen cruises. Here, we discuss the variability in DMS and DMSP_t in the north and central Indian Ocean in terms of their spatial and temporal variation. DMS and DMSP_t exhibited significant spatial and temporal variability. Apart from the concentration gradients in DMS within the Arabian Sea, Bay of Bengal and Central Indian Ocean basins, differences in average abundances were conspicuous between these basins. The Arabian Sea contained more DMS (mixed layer average was 7.8 nM) followed by the Bay of Bengal (2.8 nM) and the Central Indian Ocean (2.7 nM). The highest concentrations of DMS and DMSP_t (525 nM and 916 nM, respectively) were found in upwelling regimes along the west coast of India during the Southwest monsoon and fall intermonsoon seasons. Average surface DMS was the highest in the Arabian Sea. On the other hand observed sea-to-air fluxes of DMS were higher in the Bay of Bengal due to the prevalence of turbulent conditions. In the Arabian Sea wind speeds were low and hence the sea-to-air fluxes. The total diffusive flux of DMS from the study area to atmosphere is estimated to be about 1.02 × 10^12 g S y^−1, which contributes to 4.1–6.3% of the global DMS emission

Knowledge of the complex life cycle of Phaeocystis is a key to understanding its role in marine ecosystems and global biogeochemistry. An existing life cycle model was modified and used to integrate understanding of the Phaeocystis life cycle. In model-driven research, models expose gaps in our understanding, empirical studies ensue, and feedback improves understanding. Following this scheme, three facets of the life cycle model were examined here. With four exceptions, the empirical studies described have been presented in other literature citations. The first facet involved testing for the existence of a process or producing its description. These studies included: demonstration of in vitro colony division in Phaeocystis pouchetii , description of in vitro change in colony shape for P. pouchetii associated with senescence, determining which P. pouchetii life stage is vulnerable to viral infection and lysis, and an experiment designed to determine whether the sediment could be a source of new Phaeocystis colonies to overlying waters; results suggested that more-detailed investigation of benthic particles as a physical substrate for colony formation is warranted. The second facet involved investigation of process rate quantification or process control parameters. Process rate quantification included measurements of colony division rate and growth rate using mesocosm-derived colonies. Process control experiments included testing diatom frustule enhancement of P. pouchetii colony formation from solitary cells, and investigation of mesozooplanktonic suppression and microzooplanktonic enhancement of Phaeocystis globosa colony formation by planktonic grazer infochemicals. The third facet pertained to the molecular identification of genetic differences between single cells and colonies of P. globosa . These studies were designed to provide insight to the question of control factors involved in the transition between single cell and colonial life stages. The life cycle model provided a ready place to incorporate new insights and understanding from empirical studies into an existing model, and can be used to improve simulation models of the direct and indirect effects of Phaeocystis on global biogeochemistry.