ALBERT

Description: Denitrification was investigated in the Baltic proper at two stations with different conditions in the deep water. The Gotland Deep was examined as an example of a basin with anoxic, H2S‐containing deep water and station T was taken as an example of low‐oxygen (〈0.2 ml liter−1), sulfide‐free deep water. Denitrification was measured by the acetylene blockage method; in addition, N2O reduction was followed in samples without acetylene. To shed light on the factors limiting denitrification, we compared in situ rates to denitrification after adding nitrate or electron donors. Denitrification was restricted to the layer of the oxic‐anoxic interface in the Gotland Deep and to the water layer near the sediment of station T. For both stations it could be shown that denitrification was not limited by nitrate availability. A lack of available organic C seemed to limit denitrification rates and growth of denitrifiers. As a result of C limitation in the water column, denitrification was restricted to energy‐rich interfaces. In the low‐oxygen water away from energy‐rich interfaces, the less C‐demanding nitrification‐denitrification coupling (NH4+ → N2O → N2) seemed to be favored. Denitrification in the water of the central Baltic seems to be subjected to strong variability due to changing C supply during the course of the year. However, limitation by C availability can be assumed for most of the year and should be taken into account in calculating the N budget of the Baltic.

Description: EisenEx�the second in situ iron enrichment experiment in the Southern Ocean�was performed in the Atlantic sector over 3 weeks in November 2000 with the overarching goal to test the hypothesis that primary productivity in the Southern Ocean is limited by iron availability in the austral spring. Underwater irradiance, chlorophyll a (Chl a), photochemical efficiency, and primary productivity were measured inside and outside of an iron-enriched patch in order to quantify the response of phytoplankton to iron fertilization. Chl a concentration and photosynthetic rate (14C uptake in simulated in situ incubations) were measured in pico-, nano-, and microphytoplankton. Photochemical efficiency was studied with fast repetition rate fluorometry and xenon-pulse amplitude modulated fluorometry. The high-nutrient low-chlorophyll waters outside the Fe-enriched patch were characterized by deep euphotic zones (63-72 m), low Chl a (48-56 mg m-2), low photosynthetic efficiency (Fv/Fm ~ 0.3), and low daily primary productivity (130-220 mg C m-2 d-1). Between 70 and 90% of Chl a was found in pico- and nanophytoplankton. During the induced bloom, Fv/Fm increased up to ;0.55, primary productivity and Chl a reached the maximum values of 790 mg C m-2 d-1 and 231 mg Chl a m-2, respectively. As a consequence, the euphotic depth decreased to ~41 m. Picophytoplankton biomass hardly changed. Nano- and microphytoplankton biomass increased. In the first 2 weeks of the experiment, when the depth of the upper mixed layer was mostly 〈40 m, primary productivity was highly correlated with Chl a. In the third week, productivity was much lower than predicted from Chl a, probably because of a reduction in photosynthetic capacity as a consequence of increased physical variability in the upper water column. These results provide unequivocal evidence that iron supply is the central factor controlling phytoplankton primary productivity in the Southern Ocean, even if the mixing depth is 〉80 m.

Description: Microzooplankton have received increased attention as an important trophic link between the microbial loop and calanoid copepods. On the basis of food size spectra overlap in some microzooplankton groups and calanoid copepods, however, such microzooplankton could function as competitors rather than as food for calanoid copepods (intraguild prey). Mixotrophic flagellates presumably represent a link between the microbial loop and the micro and mesozooplankton. We investigated the effects of microzooplankton and mixotrophy by altering the presence of a heterotrophic dinoflagellate and of a mixotrophic nanoflagellate in artificial food webs with calanoid copepods as terminal consumers. Overall system productivity was manipulated by two levels of nutrient enrichment. The heterotrophic dinoflagellate drastically reduced the nanophytoplankton and enhanced the reproduction of the copepods, suggesting that its role as a competitor is negligible compared to its function as a trophic link. In spite of the presence of heterotrophic nanoflagellates, the mixotroph had a strong negative effect on the picophytoplankton and (presumably) on bacterial biomass. At the same time, the mixotroph enhanced the atomic C:N ratio of the seston biomass, indicating a higher efficiency in overall primary production. Copepod reproduction was enhanced in the presence of the mixotrophic nanoflagellate. Results did not support predictions of the intraguild predation theory: The ratios of the intraguild predators and their preys were not affected by overall system productivity

Description: Environmental evidence suggests that Aureococcus anophagefferens (Pelagophyceae), a eukaryotic picoplankton that blooms in coastal seawaters, can outcompete other organisms because of its ability to use abundant dissolved organic nitrogen (DON). To test this hypothesis, we isolated A. anophagefferens in axenic culture and monitored its growth on high-molecular weight (HMW) DON collected from sediment pore waters, a putative source for DON in bays where blooms occur. HMW DON originating from pore water had a substantially higher protein content than surface seawater DON. We found that A. anophagefferens could deplete 25-36% of the available nitrogen in cultures with HMW DON as the sole source of nitrogen and that this corresponded well with the protein fraction in pore-water HMW DON. High rates of cell surface peptide hydrolysis and no detectable N-acetyl polysaccharide hydrolysis, together with the high percentage of hydrolyzable amino acids compared to hydrolyzable aminosugars present in the HMW DON, pointed to the protein fraction as the more likely source of nitrogen used for growth. Whether or not nitrogen scavenging from protein is a common mechanism in phytoplankton is at present unknown but needs to be investigated

Description: According to a recent dynamical model, the depth of a well-mixed water column should have contrasting effects on the abundances of sinking and nonsinking phytoplankton taxa. Because of increasing light limitation, nonsinking taxa should decline monotonically with increasing mixing depth, and because of sinking loss limitation at low mixing depths, sinking taxa should peak at intermediate mixing depths. Along a gradient of mixing depths, the position of this maximum should increase with increasing taxon-specific sinking velocity and decrease with increasing background turbidity. In two field-enclosure experiments, we investigated the effects of mixing depth and background turbidity on a variety of sinking and nonsinking phytoplankton taxa. We exposed the natural, 100-µm screened phytoplankton community of a clear, unproductive, but silica-rich lake to a gradient of mixing depths (1.5- 15 m) during 4-6 weeks. To mimic two different background turbidities, the transparent enclosure walls were surrounded by either white or black foliage. Although diatoms suffered from high sedimentation losses at low mixing depths, they dominated biomass at all mixing depths throughout both experiments. Results were largely in accordance with model predictions. Specific gross growth rates of most common taxa were negatively related to mixing depth. In both experiments, the abundances of most sinking taxa showed a unimodal pattern along the mixing depth gradient, while two of three motile taxa declined monotonically with mixing depth. The depths where these taxa reached their maximal abundances were positively related to taxon-specific sinking velocity and negatively related to background turbidity.

Description: The uptake and efflux of 64Cu was studied in the marine cyanobacterium Synechoccous strain WH7803 (DC2). Uptake followed classical Michaelis-Menten type kinetics in metal-buffered seawater. The maximum uptake rate, Vmax, was 0.236 ± 0.016 × 10-18 mol Cu cell-1 h-1, with the half-saturation constant, KS, of 10-10.81±0.11 mol L-1. An efflux mechanism was also observed in WH7803, whose growth was inhibited by high internal Cu concentrations. Efflux of Cu enabled WH7803 to maintain homeostasis for Cu at typical seawater ambient free copper concentrations ([Cu2+]f). The sensitivity of WH7803 growth to Cu was related to a simple inability to regulate internal Cu concentrations when external concentrations were 〉10-11 mol L-1.

Description: Rates of cellular uptake of CO2 and HCO3- during steady-state photosynthesis were measured in the marine diatoms Thalassiosira weissflogii and Phaeodactylum tricornutum, acclimated to CO2 partial pressures of 36, 180, 360, and 1,800 ppmv. In addition, in vivo activity of extracellular (eCA) and intracellular (iCA) carbonic anhydrase was determined in relation to CO2 availability. Both species responded to diminishing CO2 supply with an increase in eCA and iCA activity. In P. tricornutum, eCA activity was close to the detection limit at higher CO2 concentrations. Simultaneous uptake of CO2 and HCO3- was observed in both diatoms. At air-equilibrated CO2 levels (360 ppmv), T. weissflogii took up CO2 and HCO3- at approximately the same rate, whereas CO2 uptake exceeded HCO3- uptake by a factor of two in P. tricornutum. In both diatoms, CO2 :HCO3- uptake ratios progressively decreased with decreasing CO2 concentration, whereas substrate affinities of CO2 and HCO3- uptake increased. Half-saturation concentrations were always 〈=5 mM CO2 for CO2 uptake and 〈700 mM HCO3- for HCO3- uptake. Our results indicate the presence of highly efficient uptake systems for CO2 and HCO3- in both diatoms at concentrations typically encountered in ocean surface waters and the ability to adjust uptake rates to a wide range of inorganic carbon supply.

Description: Redfield ratios of remineralization are calculated based on chemical data analysis on isopycnal surfaces. The concentrations of dissolved inorganic carbon used in this study were corrected for the anthropogenic CO2 content as estimated with a back-calculation technique. The corrections increased the apparent carbon remineralization by 25-30%, thus proving important for the reliable estimation of Redfield carbon ratios in the presence of anthropogenic CO2. Best estimates from this study largely confirm the more recently published Redfield ratios of remineralization. The following results were obtained for the latitude range 3-41°N along 20-29°W in the Northeast Atlantic Ocean: Corg: P ratio = 123 ± 10; Corg : N ratio = 7.2 ± 0.8; -O2 :Corg ratio = 1.34 ± 0.06; -O2 : P ratio = 165 ± 15; N: P ratio = 17.5 ± 2.0. These ratios are in close agreement with the average composition of phytoplankton and represent respiration of organic matter consisting on average of 52% protein, 36% polysaccharide, and 12% lipid.

Description: In contrast to most pelagic primary producers, benthic macrophytes pass through morphologically distinct life stages, which can be subject to different ecological controls. Using factorial field experiments, we investigated how grazing pressure (three levels) and nutrient supply (four levels) interact in controlling the passage of marine macroalgae through an apparent recruitment bottleneck at the germling stage. In comparative experiments, we asked whether relative bottom-up and top-down effects on early life stages (〈4 week germlings) vary (1) between the eutrophic Baltic Sea and the oligotrophic NW Atlantic, (2) across seasons in the NW Atlantic, and (3) among annual and perennial macroalgae. In both systems nutrient enrichment favored and grazers suppressed recruitment of green and brown annual algae; however, enrichment effects were much more pronounced in the Baltic, whereas grazer effects dominated in the NW Atlantic. Grazers induced a shift from grazer-susceptible green to more resistant brown algae in the Baltic without reducing total germling density. In the NW Atlantic, grazers strongly reduced overall recruitment rate throughout all seasons. Effects on perennials were similar in both systems with moderate losses to grazing and no effects of nutrient enrichment. Recruit densities and species composition shifted with season in the NW Atlantic. We conclude that the relative effects of grazers and nutrient enrichment depended on the nutrient status of the system, algal life history strategy, and season. Strong bottom-up and top-down controls shape benthic community composition before macroalgae reach visible size

Description: Flows of the major biogeochemical elements (C, N, P, Si) and of transparent exopolymer particles (TEP) were traced during a bloom of a natural assemblage of marine diatoms in a mesocosm (l m(3)) to determine whether the exudation and subsequent gelation of carbon-rich phytoplankton exopolymers can account for the formation and potential export of carbon in excess of that predicted by Redfield ratios. Exponential growth of the phytoplankton community in the mesocosm extended for 10 d until nitrate concentration fell below detection and concentrations of dissolved inorganic and particulate organic nitrogen and phosphorus remained stable. Tight covariation of particulate organic elements occurred as long as nutrients were replete. But, after nitrate depletion, decoupling of carbon dynamics from that of nitrogen and phosphorus was observed, with a large flow of carbon into TEP An uptake of 72% more dissolved inorganic carbon (DIC) than inferred from nitrate supply and Redfield stoichiometry (referred to as carbon overconsumption) occurred during the study, largely during the postbloom phase, and was almost entirely traced to the particulate organic matter (POM) pool. Marine snow (aggregates 〉0.5 mm) appeared at the onset of nitrate depletion and coincided with rapid increase in TEP concentrations. Elemental composition of marine snow differed from the Redfield ratio by an enrichment in carbon and a depletion in phosphorus relative to nitrogen. It is suggested that sinking of TEP-rich marine snow could be a possible mechanism for export of carbon above calculations that are based on the Redfield stoichiometry.