Description: The carbon and nitrogen content of transparent exopolymer particles (TEP) was determined and related to the concentration of TEP as quantified by a colorimetrical method. TEP were produced in the laboratory from dissolved precursors by laminar or turbulent shear. Dissolved precursors were obtained by 0.2 µm filtration from diatom cultures, with or without nutrient reduction, and from natural diatom populations. The relationship between carbon and TEP was significant, linear and species-specific. Carbon concentration of TEP derived from this relationship concurred with previous findings. Shortage of silicic acid or nitrate in the culture media had no effect on the carbon content of TEP. Molar C:N ratios of TEP were above the Redfield ratio, with a mean value of 26. It is suggested that the nitrogen fraction of TEP can be explained by adsorption of dissolved organic nitrogen (DON) onto TEP. Based on the newly established relationship, concentrations of TEP-derived carbon (TEP-C) were calculated for the Baltic Sea, the coastal Pacific, the North East Atlantic and the Northern Adriatic Sea.

Description: The abundance of transparent exopolymer particles (TEP) was determined in the northeast Atlantic Ocean (40–55°N, ∼20°W) during several cruises from June to November 1996. An accumulation of TEP in the water column was observed at bloom and post-bloom sites along a 20°W transect in June/July (maximum concentration: 124 μg Gum Xanthan equivalents (Xeq.) l−1), but concentrations were uniformly low (mean concentration: 28.5±10.2 μg Xeq. l−1) during autumn at the BIOTRANS site (47°N, 20°W). TEP concentrations in the open northeast Atlantic were considerably lower than previously published values from coastal sites. However, during June/July TEP:Chl a (weight/weight) ratios were comparable to values at coastal seas. It is suggested that phytoplankton production modulates TEP concentration in the open ocean as it does in coastal systems. TEP contributed significantly to the organic carbon pool as derived from the ratio TEP-C:POC, in summer (mean percentage: 17±7.5; w/w), as well as in autumn (mean percentage: 18±11, w/w). The potential influence of TEP on particle coagulation rates in the northeast Atlantic was assessed from estimates of their influence on particle stickiness and on particle volume concentrations. This indicated that TEP may be essential for initiating particle aggregation at low biomass concentrations, typical for open ocean sites.

Description: Incubation experiments with natural phytoplankton revealed a relationship between CO2 concentration and the production of transparent exopolymer particles (TEP), with TEP production being linearly related to theoretical CO2 uptake rates.

Description: The distribution and abundance of transparent exopolymer particles (TEP) was determined in and below pack ice of the Laptev Sea from July to September 1995. Samples were collected from the lowermost 10 cm of ice floes and at 10 cm below the ice–water interface. Abundance of bacteria, protists and TEP was determined, and the sea ice–water boundary layer was characterized using temperature, salinity and molecular viscous shear stress. TEP, with a distinct size distribution signal, were found in highest concentrations inside the sea ice, ranging from not detectable to 16 cm2 l−1 (median: 2.9 cm2 l−1). In the water, concentrations were one order of magnitude lower, ranged from below detection to 2.7 cm2 l−1 (median: 0.2 cm2 l−1) and decreased after the middle of August, whereas abundances of autotrophic flagellates (AF), diatoms, heterotrophic flagellates (HF) and ciliates increased. The abundance of TEP decreased with its size in all samples following a power law relationship. The relation of TEP to the microbial community differed between the sea ice and water, being positively correlated with bacteria and diatoms in the ice and negatively correlated with HF in the sea water. The presence of a pycnocline significantly influenced the abundance of organisms, diatom composition and TEP concentrations. Pennate diatoms dominated by Nitzschia frigida were most abundant inside the ice. Though bacteria have the potential to produce exopolymeric substances (EPS), the results of this study indicate that the majority of TEP at the ice–water interface in first-year Arctic summer pack ice are produced by diatoms.

Description: The termination of diatom spring blooms in temperate waters has been connected with the formation and subsequent rapid sedimentation of aggregates. According to coagulation theory, the rate of aggregate formation depends on the probability of particle collision and on the efficiency with which two particles adhere once they have collided (stickiness). During this study, the variation in particle stickiness was determined over the decline of a diatom bloom using the Couette Chamber assay with low shear (G = 0.86 s–1). A mixed diatom population, dominated by Skeletonema costatum, was sampled during the spring bloom in the Baltic Sea and incubated in the laboratory for 18 days. Measurements of diatom species composition, transparent exopolymer particles (TEP) and bulk particle abundance, as well as chemical and biological variables, were conducted in order to reveal the determinants of coagulation efficiency. The investigation showed that an increase in TEP concentration relative to conventional particles at the decline of the bloom significantly enhanced apparent coagulation efficiencies. High proportions of TEP led to apparent values of stickiness 〉1, which indicates that collision rates can be substantially underestimated when the stickiness parameter α is calculated on the basis of conventional particle counting only, e.g. with the Coulter Counter. A new stickiness parameter, α′, was therefore estimated based on the combined volume fractions of TEP and conventional particles. The problems of stickiness measurements are discussed and the role of TEP in coagulation processes is emphasized.

Description: A simple two-size-class aggregation model is developed to describe the time-dependent carbon content of dissolved polysaccharides (PCHO) and of transparent exopolymer particles (TEP) during the bloom. A conservative estimate for the effective collision kernel is obtained from the Smoluchowski equation under the assumption that the growth of aggregates is controlled by a Brownian process near the scaling regime. In the model, PCHO are assumed to represent a fraction of the photosynthetic carbon, which is not used for net algal growth. Time dependence of chlorophyll a and of cellular carbon during the bloom is modelled in terms of algal growth and sinking of single and aggregated algal cells. The aggregation of exopolysaccharides into TEP may have important implications for the organic carbon cycle in the ocean, as TEP promote the aggregation of algae during a bloom.

Description: One particular task of marine ecosystem models is to simulate the biogenic transformation of dissolved inorganic carbon (DIC) into organic matter and hence to quantify the export of particulate organic carbon (POC) to deep oceanic layers. To date, environmental changes, such as increasing carbon dioxide concentrations (pCO_2) and temperature, are perceived to have an impact on the formation of organic carbon. However, well established nitrogen or phosphorus based ecosystem models are insensitive to variations in the carbonate system. In order to investigate biological responses to pCO_2 variations, ecosystem models need to distinguish between carbon, nitrogen, and/or phosphorus cycles. We present a simple biological model which decouples carbon from nitrogen fluxes such that carbon found in transparent exopolymer particles (TEP) is additionally accounted for. The model regards phytoplankton acclimation to varying environmental conditions, having included parameterizations for phytoplankton growth as proposed by Geider et al.~(1998, L&O). By means of data assimilation, an optimal parameter set is determined, which brings model results into agreement with experimental data. From the optimised model results it is infered that about 50% of dissolved organic carbon (DOC) exuded by phytoplankton is subsequently transformed into TEP, eventually influencing the amount of POC available for the export flux. Model sensitivity studies are performed at local sites and along a latitudinal transect (30^oN-60^oN at 19^oW) in the North Atlantic. As soon as CO_2 limitation for phytoplankton growth is explicitely considered in the model, the formation of POC shows great sensitivity to pCO_2 variations. Temperature variations alter remineralisation rates and growth efficiencies. With the current model version dependencies between biomass accumulation, the date of nutrient depletion to occur, and the exudation of organic compounds are acquired.

Description: The export of organic carbon to the deep ocean is mediated by sinking of large particles, such as marine snow, the formation of which is enhanced in the presence of transparent exopolymer particles (TEP) . TEP form from dissolved and colloidal polysaccharides by aggregation processes. Especially when running into nutrient limitation phytoplankton organisms are a source of TEP in pelagic ecosystems as the cells release a significant amount of the assimilated carbon in the form of polysaccharides. Because CO_2 concentration influences carbon assimilation rates, we hypothesized that polysaccharide exudation and aggregation into TEP is related to CO_2 concentration under nutrient limiting conditions. We tested this hypothesis in several lab and outdoor experiments with natural populations and cultures of phytoplankton exposed to various levels of CO_2 concentrations. Our results indicate that TEP production increases with CO_2 concentration and provides an enhanced sink for carbon during phytoplankton blooms.

Description: A fraction of the photosynthetically fixed carbon is not used for phytoplankton growth, but channelled to the outer medium via exudation. This fraction include carbon rich exopolymers that coagulate to particles, such as transparent exopolymer particles (TEP). Through aggregation with cells and debris, TEP are incorporated into large, rapidly settling marine snow and contribute to the vertical flux of organic matter to the deep sea. The influence of TEP formation on the C:N:P stoichiometry of marine phy- toplankton blooms was examined during two mesocosm studies. During the blooms, which were dominated by a natural assembly of marine diatoms and the calcifying coccolithophorid Emiliania huxleyi, respectively, an increase of TEP concentration was observed immediately after nutrient depletion, followed by the appearance of ma- rine snow. The contribution of TEP to the carbon flow during both blooms indicates that sinking of TEP-rich marine snow is a possible mechanism for the removal of carbon from surface waters above calculations based on Redfield stoichiometry.