Description: The distribution of transparent exopolymer particles (TEP) was investigated during a coccolithophorid bloom in the northern Bay of Biscay (North Atlantic Ocean) in early June 2006. MODIS chlorophyll-a (Chl-a) and reflectance images before and during the cruise were used to localize areas of important biological activity and high reflectance (HR). TEP profiles along the continental margin, determined using microscopic (TEPmicro) and colorimetric (TEPcolor) methods, showed abundant (6.1×106–4.4×107 L−1) and relatively small (0.5–20 μm) particles, leading to a low total volume fraction (0.05–2.2 ppm) of TEPmicro and similar vertical profiles of TEPcolor. Estimates of carbon content in TEP (TEP-C) derived from the microscopic approach yielded surface concentration of 1.50 μmol C L−1. The contribution of TEP-C to particulate organic carbon (POC) was estimated to be 12% (molar C ratio) during this survey. Our results suggest that TEP formation is a probable first step to rapid and efficient export of C during declining coccolithophorid blooms.

Description: Observations that the majority of silica dissolution occurs within the upper 200 m of the ocean, and that sedimentation rates of diatom frustules generally do not decrease significantly with depth, suggested reduced dissolution rates of diatoms embedded within sinking aggregates. To investigate this hypothesis, silica dissolution rates of aggregated diatom cells were compared to those of dispersed cells during conditions mimicking sedimentation below the euphotic zone. Changes in the concentrations of biogenic silica, silicic acid, cell numbers, chlorophyll a and transparent exopolymer particles (TEP) were monitored within aggregates and in the surrounding seawater (SSW) during two 42-day experiments. Whereas the concentration of dispersed diatoms decreased over the course of the experiment, the amount of aggregated cells remained roughly constant after an initial increase. Initially only 6% of cells were aggregated and at the end of the experiment more than 60% of cells were enclosed within aggregates. These data imply lower dissolution rates for aggregated cells. However, fluxes of silica between the different pools could not be constrained reliably enough to unequivocally prove reduced dissolution for aggregated cells.

Description: The present study investigates the combined effect of phosphorous limitation, elevated partial pressure of CO2 (pCO2) and temperature on a calcifying strain of Emiliania huxleyi (PML B92/11) by means of a fully controlled continuous culture facility. Two levels of phosphorous limitation were consecutively applied by renewal of culture media (N:P=26) at dilution rates (D) of 0.3 d−1 and 0.1 d−1. CO2 and temperature conditions were 300, 550 and 900 μatm pCO2 at 14 °C and 900 μatm pCO2 at 18 °C. In general, the steady state cell density and particulate organic carbon (POC) production increased with pCO2, yielding significantly higher concentrations in cultures grown at 900 μatm pCO2 compared to 300 and 550 μatm pCO2. At 900 μatm pCO2, elevation of temperature as expected for a greenhouse ocean, further increased cell densities and POC concentrations. In contrast to POC concentration, C-quotas (pmol C cell−1) were similar at D=0.3 d−1 in all cultures. At D=0.1 d−1, a reduction of C-quotas by up to 15% was observed in the 900 μatm pCO2 at 18 °C culture. As a result of growth rate reduction, POC:PON:POP ratios deviated strongly from the Redfield ratio, primarily due to an increase in POC. Ratios of particulate inorganic and organic carbon (PIC:POC) ranged from 0.14 to 0.18 at D=0.3 d−1, and from 0.11 to 0.17 at D=0.1 d−1, with variations primarily induced by the changes in POC. At D=0.1 d−1, cell volume was reduced by up to 22% in cultures grown at 900 μatm pCO2. Our results indicate that changes in pCO2, temperature and phosphorus supply affect cell density, POC concentration and size of E. huxleyi (PML B92/11) to varying degrees, and will likely impact bloom development as well as biogeochemical cycling in a greenhouse ocean.

Description: Primary production (PP), calcification (CAL), bacterial production (BP) and dark community respiration (DCR) were measured along with a set of various biogeochemical variables, in early June 2006, at several stations at the shelf break of the northern Bay of Biscay. The cruise was carried out after the main spring diatom bloom that, based on the analysis of a time-series of remotely sensed chlorophyll-a (Chl-a), peaked in mid-April. Remotely sensed sea surface temperature (SST) indicated the occurrence of enhanced vertical mixing (due to internal tides) at the continental slope, while adjacent waters on the continental shelf were stratified, as confirmed by vertical profiles of temperature acquired during the cruise. The surface layer of the stratified water masses (on the continental shelf) was depleted of inorganic nutrients. Dissolved silicate (DSi) levels probably did not allow significant diatom development. We hypothesize that mixing at the continental slope allowed the injection of inorganic nutrients that triggered the blooming of mixed phytoplanktonic communities dominated by coccolithophores (Emiliania huxleyi) that were favoured with regards to diatoms due to the low DSi levels. Based on this conceptual frame, we used an indicator of vertical stratification to classify the different sampled stations, and to reconstruct the possible evolution of the bloom from the onset at the continental slope (triggered by vertical mixing) through its development as the water mass was advected on-shelf and stratified. We also established a carbon mass balance at each station by integrating in the photic layer PP, CAL and DCR. This allowed computation at each station of the contribution of PP, CAL and DCR to CO2 fluxes in the photic layer, and how they changed from one station to another along the sequence of bloom development (as traced by the stratification indicator). This also showed a shift from net autotrophy to net heterotrophy as the water mass aged (stratified), and suggested the importance of extracellular production of carbon to sustain the bacterial demand in the photic and aphotic layers.

Description: To investigate the role of ballasting by biogenic minerals in the export of organic matter in the ocean, a laboratory experiment was conducted comparing aggregate formation and settling velocity of non-calcifying and calcifying strains of the coccolithophore Emiliania huxleyi. Experiments were conducted by making aggregates using a roller table and following aggregate properties during incubation for a period of 40 days. Size, shape, and settling velocities of aggregates were described by image analysis of video pictures recorded during the roller tank incubation. Our results show that biogenic calcite has a strong effect on the formation rate and abundance of aggregates and on aggregate properties such as size, excess density, porosity, and settling velocity. Aggregates of calcifying cells (AGGCAL) formed faster, were smaller and had higher settling velocities, excess densities, and mass than those of non-calcifying cells (AGGNCAL). AGGCAL showed no loss during the duration of the experiment, whereas AGGNCAL decreased in size after 1 month of incubation. Potential mechanisms that can explain the different patterns in aggregate formation are discussed. Comparison of settling velocities of AGGCAL and AGGNCAL with aggregates formed by diatoms furthermore indicated that the ballast effect of calcite is greater than that of opal. Together these results help to better understand why calcite is of major importance for organic matter fluxes to the deep ocean.

Description: Protecting the ocean has become a major goal of international policy as human activities increasingly endanger the integrity of the ocean ecosystem, often summarized as “ocean health.” By and large, efforts to protect the ocean have failed because, among other things, (1) the underlying socio-ecological pathways have not been properly considered, and (2) the concept of ocean health has been ill defined. Collectively, this prevents an adequate societal response as to how ocean ecosystems and their vital functions for human societies can be protected and restored. We review the confusion surrounding the term “ocean health” and suggest an operational ocean-health framework in line with the concept of strong sustainability. Given the accelerating degeneration of marine ecosystems, the restoration of regional ocean health will be of increasing importance. Our advocated transdisciplinary and multi-actor framework can help to advance the implementation of more active measures to restore ocean health and safeguard human health and well-being.

Description: We investigated the effect of CO2 and primary production on the carbon isotopic fractionation of alkenones and particulate organic matter (POC) during a natural phytoplankton bloom dominated by the coccolithophore Emiliania huxleyi. In nine semi-closed mesocosms (~11 m3 each), three different CO2 partial pressures (pCO2) in triplicate represented glacial (~180 ppmv CO2), present (~380 ppmv CO2), and year 2100 (~710 ppmv CO2) CO2 conditions. The largest shift in alkenone isotopic composition (4–5&) occurred during the exponential growth phase, regardless of the CO2 concentration in the respective treatment. Despite the difference of ~500 ppmv, the influence of pCO2 on isotopic fractionation was marginal (1–2&). During the stationary phase, E. huxleyi continued to produce alkenones, accumulating cellular concentrations almost four times higher than those of exponentially dividing cells. Our isotope data indicate that, while alkenone production was maintained, the interaction of carbon source and cellular uptake dynamics by E. huxleyi reached a steady state. During stationary phase, we further observed a remarkable increase in the difference between d13C of bulk organic matter and of alkenones spanning 7–12&. We suggest that this phenomenon is caused mainly by a combination of extracellular release of 13C-enriched polysaccharides and subsequent particle aggregation induced by the production of transparent exopolymer particles (TEP). 2007 Elsevier Inc. All rights reserved.

Description: Suspended particles and particle aggregates, which formed from concentrated field samples on the roller table, were characterized biologically and chemically along a transect through the Baltic Sea in summer 1999. Phytoplankton composition in field samples was dominated by cyanobacteria, including the filamentous diazotrophic cyanobacteria Aphanizomenon ‘ baltica’, Nodularia spumigena and Anabaena spp. These species formed aggregates together with diatoms, mainly Skeletonema costatum and Chaetoceros spp. and with dinoflagellates, mainly withDinophysis norvegica . Compared to the Redfield ratio, concentration ratios of particulate organic carbon, nitrogen and phosphorus, [POC]:[PON]:[POP], indicated an enrichment of carbon, especially in aggregates. However, regression analysis indicated a higher production rate of PON relative to POP and POC and significant background concentrations of POC. In field samples the concentration of transparent exopolymer particles (TEP) varied around 200 μg Xanthan Equiv. l−1 and comprised a volume fraction of 2–7 ppm and an abundance of about 105 TEP ml−1. TEP were enriched in aggregates as inferred from volume ratios of TEP to conventional particles. It is suggested, that TEP contribute substantially to the background concentration of POC, while the high production rate of PON is attributed to nitrogen fixation of diazotrophic cyanobacteria.

Description: Highlights: • The epoch of the Anthropocene, a period during which human activity has been the dominant influence on climate and the environment, has witnessed a decline in oxygen concentrations and an expansion of oxygen-depleted environments in both coastal and open ocean systems since the middle of the 20th century. • This review paper provides a synthesis of system-specific drivers of low oxygen in a range of case studies representing marine systems in the open ocean, on continental shelves, in enclosed seas and in the coastal environment. • Identification of similar and contrasting responses within and across system types and corresponding oxygen regimes is shown to be informative both in understanding and isolating key controlling processes and provides a sound basis for predicting change under anticipated future conditions. • Case studies were selected to achieve a balance in system diversity and global coverage. • Each case study describes system attributes, including the present-day oxygen environment and known trends in oxygen concentrations over time. • Central to each case study is the identification of the physical and biogeochemical processes that determine oxygen concentrations through the tradeoff between ventilation and respiration. • Spatial distributions of oxygen and time series of oxygen data provide the opportunity to identify trends in oxygen availability and have allowed various drivers of low oxygen to be distinguished through correlative and causative relationships. • Deoxygenation results from a complex interplay of hydrographic and biogeochemical processes and the superposition of these processes, some additive and others subtractive, makes attribution to any particular driver challenging. • System-specific models are therefore required to achieve a quantitative understanding of these processes and of the feedbacks between processes at varying scales. Abstract: The epoch of the Anthropocene, a period during which human activity has been the dominant influence on climate and the environment, has witnessed a decline in oxygen concentrations and an expansion of oxygen-depleted environments in both coastal and open ocean systems since the middle of the 20th century. This paper provides a review of system-specific drivers of low oxygen in a range of case studies representing marine systems in the open ocean, on continental shelves, in enclosed seas and in the coastal environment. Identification of similar and contrasting responses within and across system types and corresponding oxygen regimes is shown to be informative both in understanding and isolating key controlling processes and provides a sound basis for predicting change under anticipated future conditions. Case studies were selected to achieve a balance in system diversity and global coverage. Each case study describes system attributes, including the present-day oxygen environment and known trends in oxygen concentrations over time. Central to each case study is the identification of the physical and biogeochemical processes that determine oxygen concentrations through the tradeoff between ventilation and respiration. Spatial distributions of oxygen and time series of oxygen data provide the opportunity to identify trends in oxygen availability and have allowed various drivers of low oxygen to be distinguished through correlative and causative relationships. Deoxygenation results from a complex interplay of hydrographic and biogeochemical processes and the superposition of these processes, some additive and others subtractive, makes attribution to any particular driver challenging. System-specific models are therefore required to achieve a quantitative understanding of these processes and of the feedbacks between processes at varying scales.

Description: Highlights: • Adriatic coastal area: simultaneous atmospheric and seawater field measurements • Nutrient variability in aerosols, rainwater and atmospheric deposition (AD) fluxes • Local open-fire biomass burning (BB) affected chemistry of atmospheric samples. • AD impacted nutrient levels and N:P ratios in the sea surface microlayer (SML). • The SML plankton development and organic matter enrichments followed BB episodes. Abstract: Atmospheric deposition (AD) of nutrients and its impact on the sea surface requires consideration of interfacial processes within the sea surface microlayer (SML), the ocean-atmosphere boundary layer of major importance for many global biogeochemical and climate-related processes. This study comprised a comprehensive dataset, including dissolved NO3−, NH4+ and PO43− in ambient aerosol particles, wet deposition and sea surface samples collected from February to July 2019 at a central Adriatic coastal site. The aerosol mean concentration of dissolved nitrogen (DIN = NO3− + NH4+) and PO43− were 48.8 ± 82.8 μmol m−3 and 0.8 ± 0.6 μmol m−3, respectively, while their total fluxes (dry + wet) ranged from 24.2 to 212.3 μmol m−2 d−1 (mean 123.2 ± 53.2 μmol m−2 d−1) and from 1.2 to 2.1 μmol m−2 d−1 (mean 1.5 ± 0.3 μmol m−2 d−1), respectively. Intensive local episodes of open biomass burning (BB) significantly increased aerosol DIN concentrations as well as DIN deposition fluxes, particularly altering the molar DIN/PO43− ratio of atmospheric samples. The DIN temporal patterns showed high variability in the SML (range 0.2–24.6 μmol L−1, mean 5.0 ± 7.1 μmol L−1) in contrast to the underlying water samples (range 0.5–4.2 μmol L−1, mean 1.9 ± 1.2 μmol L−1), with significant increases during BB periods. Variability in abundance of heterotrophic bacteria and autotrophs in the SML along with concentrations of bulk dissolved and particulate organic carbon as well as dissolved and particulate lipids and carbohydrates, gel particles and surfactants followed DIN enhancements with a two-week delay. This study showed that AD can affect the short-term scale enrichments of organic matter in the SML, especially when accompanied by BB emissions typical of the overall Mediterranean coastal environment. This could have strong implications for global air-sea exchange processes, including those of climate relevant gases, mediated by the SML.