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The science guiding the EUREC4A campaign and its measurements is presented. EUREC4A comprised roughly 5 weeks of measurements in the downstream winter trades of the North Atlantic - eastward and southeastward of Barbados. Through its ability to characterize processes operating across a wide range of scales, EUREC4A marked a turning point in our ability to observationally study factors influencing clouds in the trades, how they will respond to warming, and their link to other components of the earth system, such as upper-ocean processes or the life cycle of particulate matter. This characterization was made possible by thousands (2500) of sondes distributed to measure circulations on meso- (200 km) and larger (500 km) scales, roughly 400 h of flight time by four heavily instrumented research aircraft; four global-class research vessels; an advanced ground-based cloud observatory; scores of autonomous observing platforms operating in the upper ocean (nearly 10 000 profiles), lower atmosphere (continuous profiling), and along the air-sea interface; a network of water stable isotopologue measurements; targeted tasking of satellite remote sensing; and modeling with a new generation of weather and climate models. In addition to providing an outline of the novel measurements and their composition into a unified and coordinated campaign, the six distinct scientific facets that EUREC4A explored - from North Brazil Current rings to turbulence-induced clustering of cloud droplets and its influence on warm-rain formation - are presented along with an overview of EUREC4A's outreach activities, environmental impact, and guidelines for scientific practice. Track data for all platforms are standardized and accessible at 10.25326/165 , and a film documenting the campaign is provided as a video supplement.

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Recent modeling results suggest that oceanic oxygen levels will decrease significantly over the next decades to centuries in response to climate change and altered ocean circulation. Hence the future ocean may experience major shifts in nutrient cycling triggered by the expansion and intensification of tropical oxygen minimum zones (OMZs). There are numerous feedbacks between oxygen concentrations, nutrient cycling and biological productivity; however, existing knowledge is insufficient to understand physical, chemical and biological interactions in order to adequately assess past and potential future changes. We investigated the pelagic biogeochemistry of OMZs in the eastern tropical North Atlantic and eastern tropical South Pacific during a series of cruise expeditions and mesocosm studies. The following summarizes the current state of research on the influence of low environmental oxygen conditions on marine biota, viruses, organic matter formation and remineralization with a particular focus on the nitrogen cycle in OMZ regions. The impact of sulfidic events on water column biogeochemistry, originating from a specific microbial community capable of highly efficient carbon fixation, nitrogen turnover and N2O production is further discussed. Based on our findings, an important role of sinking particulate organic matter in controlling the nutrient stochiometry of the water column is suggested. These particles can enhance degradation processes in OMZ waters by acting as microniches, with sharp gradients enabling different processes to happen in close vicinity, thus altering the interpretation of oxic and anoxic environments.

Aphanizomenon is a widespread genus of nitrogen (N2)-fixing cyanobacteria in lakes and estuaries, accounting for a large fraction of the summer N2-fixation in the Baltic Sea. However, information about its cell-specific carbon (C)- and N2-fixation rates in the early growth season has not previously been reported. We combined various methods to study N2-fixation, photosynthesis and respiration in field-sampled Baltic Sea Aphanizomenon sp. during early summer at 10°C. Stable isotope incubations at in situ light intensities during 24 h combined with cell-specific secondary ion mass spectrometry showed an average net N2-fixation rate of 55 fmol N cell(-1) day(-1). Dark net N2-fixation rates over a course of 12 h were 20% of those measured in light. C-fixation, but not N2-fixation, was inhibited by high ambient light intensities during daytime. Consequently, the C:N fixation ratio varied substantially over the diel cycle. C- and N2-fixation rates were comparable to those reported for Aphanizomenon sp. in August at 19°C, using the same methods. High respiration rates (23% of gross photosynthesis) were measured with (14)C-incubations and O2-microsensors, and presumably reflect the energy needed for high N2-fixation rates. Hence, Aphanizomenon sp. is an important contributor to N2-fixation at low in situ temperatures in the early growth season.

Aphanizomenon is a widespread genus of nitrogen (N2)-fixing cyanobacteria in lakes and estuaries, accounting for a large fraction of the summer N2-fixation in the Baltic Sea. However, information about its cell-specific carbon (C)- and N2-fixation rates in the early growth season has not previously been reported. We combined various methods to study N2-fixation, photosynthesis and respiration in field-sampled Baltic Sea Aphanizomenon sp. during early summer at 10°C. Stable isotope incubations at in situ light intensities during 24 h combined with cell-specific secondary ion mass spectrometry showed an average net N2-fixation rate of 55 fmol N cell−1 day−1. Dark net N2-fixation rates over a course of 12 h were 20% of those measured in light. C-fixation, but not N2-fixation, was inhibited by high ambient light intensities during daytime. Consequently, the C:N fixation ratio varied substantially over the diel cycle. C- and N2-fixation rates were comparable to those reported for Aphanizomenon sp. in August at 19°C, using the same methods. High respiration rates (23% of gross photosynthesis) were measured with 14C-incubations and O2-microsensors, and presumably reflect the energy needed for high N2-fixation rates. Hence, Aphanizomenon sp. is an important contributor to N2-fixation at low in situ temperatures in the early growth season.

The release of organic material upon algal cell lyses has a key role in structuring bacterial communities and affects the cycling of biolimiting elements in the marine environment. Here we show that already before cell lysis the leakage or excretion of organic matter by infected yet intact algal cells shaped North Sea bacterial community composition and enhanced bacterial substrate assimilation. Infected algal cultures of Phaeocystis globosa grown in coastal North Sea water contained gamma-and alphaproteobacterial phylotypes that were distinct from those in the non-infected control cultures 5 h after infection. The gammaproteobacterial population at this time mainly consisted of Alteromonas sp. cells that were attached to the infected but still intact host cells. Nano-scale secondary-ion mass spectrometry (nanoSIMS) showed similar to 20% transfer of organic matter derived from the infected C-13- and N-15-labelled P. globosa cells to Alteromonas sp. cells. Subsequent, viral lysis of P. globosa resulted in the formation of aggregates that were densely colonised by bacteria. Aggregate dissolution was observed after 2 days, which we attribute to bacteriophage-induced lysis of the attached bacteria. Isotope mass spectrometry analysis showed that 40% of the particulate C-13-organic carbon from the infected P. globosa culture was remineralized to dissolved inorganic carbon after 7 days. These findings reveal a novel role of viruses in the leakage or excretion of algal biomass upon infection, which provides an additional ecological niche for specific bacterial populations and potentially redirects carbon availability.

Eastern boundary upwelling systems (EBUS) are regions of high primary production often associated with oxygen minimum zones (OMZs). They represent key regions for the oceanic nitrogen (N) cycle. By exporting organic matter (OM) and nutrients produced in the coastal region to the open ocean, EBUS can play an important role in sustaining primary production in subtropical gyres. However, losses of fixed inorganic N through denitrification and anammox processes take place in oxygen depleted environments such as EBUS, and can potentially mitigate the role of these regions as a source of N to the open ocean. EBUS can also represent a considerable source of nitrous oxide (N2O) to the atmosphere, affecting the atmospheric budget of N2O. In this paper a 3-D coupled physical/biogeochemical model (ROMS/BioEBUS) is used to investigate the N budget in the Namibian upwelling system. The main processes linked to EBUS and associated OMZs are taken into account. The study focuses on the northern part of the Benguela upwelling system (BUS), especially the Walvis Bay area (between 22 degrees S and 24 degrees S) where the OMZ is well developed. Fluxes of N off the Walvis Bay area are estimated in order to understand and quantify (1) the total N offshore export from the upwelling area, representing a possible N source that sustains primary production in the South Atlantic subtropical gyre; (2) export production and subsequent losses of fixed N via denitrification and anammox under suboxic conditions (O-2 < 25 mmol O-2 m(-3)); and (3) the N2O emission to the atmosphere in the upwelling area. In the mixed layer, the total N offshore export is estimated as 8.5+/-3.9x10(10) mol N yr(-1) at 10 degrees E off the Walvis Bay area, with a mesoscale contribution of 20 %. Extrapolated to the whole BUS, the coastal N source for the subtropical gyre corresponds to 0.1+/-0.04 mol N m(-2) yr(-1). This N flux represents a major source of N for the gyre compared with other N sources, and contributes 28%

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Phaeocystis globosa is an ecologically important bloom-forming phytoplankton, which sequesters substantial amounts of inorganic carbon and can form carbon-enriched chitinous star-like structures. Viruses infecting P.globosa (PgVs) play a significant regulatory role in population dynamics of the host species. However, the extent to which viruses alter host physiology and its carbon assimilation on single cell level is still largely unknown. This study demonstrates for the first time the impact of viral infection on carbon assimilation and cell morphology of individual axenic P.globosa cells using two single cell techniques: high resolution nanometre-scale Secondary-Ion Mass Spectrometry (nanoSIMS) approach and atomic force microscopy (AFM). Up until viral lysis (19h post infection), the bulk carbon assimilation by infected P.globosa cultures was identical to the assimilation by the non-infected cultures (33 mu molCl1). However, single cell analysis showed that viral infection of P.globosa impedes the release of star-like structures. Non-infected cells transfer up to 44.5 mu molCl1 (36%) of cellular biomass in the form of star-like structures, suggesting a vital role in the survival of P.globosa cells. We hypothesize that impediment of star-like structures in infected P.globosa cells may inactivate viral infectivity by forming flocculants after cell lysis. Moreover, we show that substantial amounts of newly produced viruses (approximate to 68%) were attached to P.globosa cells prior to cell lysis. Further, we speculate that infected cells become more susceptible for grazing which provides potential reasons for the sudden disappearance of PgVs in the environment. The scenarios of enhanced grazing is at odds to the current perspective that viral infections facilitates microbial mediated processes by diverting host material away from the higher trophic levels.

The Eastern Boundary Upwelling Systems (EBUS) contribute to one fifth of the global catches in the ocean. Often associated with Oxygen Minimum Zones (OMZs), EBUS represent key regions for the oceanic nitrogen (N) cycle. Important bioavailable N loss due to denitrification and anammox processes as well as greenhouse gas emissions (e.g, N2O) occur also in these EBUS. However, their dynamics are currently crudely represented in global models. In the climate change context, improving our capability to properly represent these areas is crucial due to anticipated changes in the winds, productivity, and oxygen content. We developed a biogeochemical model (BioEBUS) taking into account the main processes linked with EBUS and associated OMZs. We implemented this model in a 3-D realistic coupled physical/biogeochemical configuration in the Namibian upwelling system (northern Benguela) using the high-resolution hydrodynamic ROMS model. We present here a validation using in situ and satellite data as well as diagnostic metrics and sensitivity analyses of key parameters and N2O parameterizations. The impact of parameter values on the OMZ off Namibia, on N loss, and on N2O concentrations and emissions is detailed. The model realistically reproduces the vertical distribution and seasonal cycle of observed oxygen, nitrate, and chlorophyll a concentrations, and the rates of microbial processes (e.g, NH4+ and NO2− oxidation, NO3− reduction, and anammox) as well. Based on our sensitivity analyses, biogeochemical parameter values associated with organic matter decomposition, vertical sinking, and nitrification play a key role for the low-oxygen water content, N loss, and N2O concentrations in the OMZ. Moreover, the explicit parameterization of both steps of nitrification, ammonium oxidation to nitrate with nitrite as an explicit intermediate, is necessary to improve the representation of microbial activity linked with the OMZ. The simulated minimum oxygen concentrations are driven by th

Phaeocystis globosa is an ecologically important bloom-forming phytoplankton, which sequesters substantial amounts of inorganic carbon and can form carbon-enriched chitinous star-like structures. Viruses infecting P. globosa (PgVs) play a significant regulatory role in population dynamics of the host species. However, the extent to which viruses alter host physiology and its carbon assimilation on single cell level is still largely unknown. This study demonstrates for the first time the impact of viral infection on carbon assimilation and cell morphology of individual axenic P. globosa cells using two single cell techniques: high resolution nanometre-scale Secondary-Ion Mass Spectrometry (nanoSIMS) approach and atomic force microscopy (AFM). Up until viral lysis (19 h post infection), the bulk carbon assimilation by infected P. globosa cultures was identical to the assimilation by the non-infected cultures (33 micromol C l(-1) ). However, single cell analysis showed that viral infection of P. globosa impedes the release of star-like structures. Non-infected cells transfer up to 44.5 micromol C l(-1) (36%) of cellular biomass in the form of star-like structures, suggesting a vital role in the survival of P. globosa cells. We hypothesize that impediment of star-like structures in infected P. globosa cells may inactivate viral infectivity by forming flocculants after cell lysis. Moreover, we show that substantial amounts of newly produced viruses ( approximately 68%) were attached to P. globosa cells prior to cell lysis. Further, we speculate that infected cells become more susceptible for grazing which provides potential reasons for the sudden disappearance of PgVs in the environment. The scenarios of enhanced grazing is at odds to the current perspective that viral infections facilitates microbial mediated processes by diverting host material away from the higher trophic levels.

FS Meteor Fahrt 93 Wochenbericht 4 (25.02.-03.03.2013)

The release of organic material upon algal cell lyses has a key role in structuring bacterial communities and affects the cycling of biolimiting elements in the marine environment. Here we show that already before cell lysis the leakage or excretion of organic matter by infected yet intact algal cells shaped North Sea bacterial community composition and enhanced bacterial substrate assimilation. Infected algal cultures of Phaeocystis globosa grown in coastal North Sea water contained gamma- and alphaproteobacterial phylotypes that were distinct from those in the non-infected control cultures 5 h after infection. The gammaproteobacterial population at this time mainly consisted of Alteromonas sp. cells that were attached to the infected but still intact host cells. Nano-scale secondary-ion mass spectrometry (nanoSIMS) showed ~20% transfer of organic matter derived from the infected 13C- and 15N-labelled P. globosa cells to Alteromonas sp. cells. Subsequent, viral lysis of P. globosa resulted in the formation of aggregates that were densely colonised by bacteria. Aggregate dissolution was observed after 2 days, which we attribute to bacteriophage-induced lysis of the attached bacteria. Isotope mass spectrometry analysis showed that 40% of the particulate 13C-organic carbon from the infected P. globosa culture was remineralized to dissolved inorganic carbon after 7 days. These findings reveal a novel role of viruses in the leakage or excretion of algal biomass upon infection, which provides an additional ecological niche for specific bacterial populations and potentially redirects carbon availability.

Fixed nitrogen (N) loss to biogenic N2 in intense oceanic O2 minimum zones (OMZ) accounts for a large fraction of the global N sink and is an essential control on the ocean's N budget. However, major uncertainties exist regarding microbial pathways as well as net impact on the magnitude of N-loss and the ocean's overall N budget. Here we report the discovery of a N-loss hotspot in the Peru OMZ associated with a coastally trapped mesoscale eddy that is marked by an extreme N deficit matched by biogenic N2 production, high NO2− levels, and the highest isotope enrichments observed so far in OMZ's for the residual NO3−. High sea surface chlorophyll (SSC) in seaward flowing streamers provides evidence for offshore eddy transport of highly productive, inshore water. Resulting pulses in the downward flux of particles likely stimulated heterotrophic dissimilatory NO3− reduction and subsequent production of biogenic N2. The associated temporal/spatial heterogeneity of N-loss, mediated by a local succession of microbial processes, may explain inconsistencies observed among prior studies. Similar transient enhancements of N-loss likely occur within all other major OMZ's exerting a major influence on global ocean N and N isotope budgets.

ID: 10044 Microbially-mediated N-loss in intense oceanic O2 minimum zones (OMZ) accounts for a large fraction of the global sink and is an essential control on the ocean’s N budget. However, major uncertainties exist regarding pathways and their net impact on N-loss magnitude. To address these issues, relevant biogeochemical properties were mapped throughout the Peru OMZ in winter 2009. South of 7 to 10°S, low O2 concentrations in the Peru OMZ enabled N-loss processes as shown by the appearance of nitrite, net removal of nitrate from the water column, and isotopic enrichment in the residual nitrate. However, two stations exhibited uniquely extreme values. Satellite imagery shows these stations to be associated with the seaward edges of coastal mesoscale eddies. We hypothesize that eddies transport productive, nutrient-rich coastal water offshore that create deep-ocean N-loss ‘hotspots’ with extreme N isotope fractionation. As a result, eddies are likely important for overall enhancement of N-loss from the Peru OMZ. Similar transient enhancements of N-loss by eddies likely occur within all other major OMZ’s thereby exerting a major influence on global ocean N and N isotope budgets.

Foraminifera are an important faunal element of the abyssal ecosystem and largely depend on deposited particulate organic matter from the photic zone to sustain their metabolism for growth and reproduction. However, their role in the carbon cycle in deep-sea sediments is insufficiently studied. We investigated benthic foraminifera at Station M (4000 m depth) in the Northeast Pacific and assessed the response of individual species to a simulated phytodetritus pulse during an in situ feeding experiment. Sediments were incubated for 4 d with 13C-labeled diatoms (Thalassiosira weissflogii) applied to the sediment surface. The living foraminiferal community (>0.063 mm) of the upper 3 cm contained >100 species and was strongly dominated by a few taxa of soft-walled saccamminids. Population density of the entire living foraminiferal community was highest at the sediment surface (mean ± SD = 279 ± 72 ind. 10 cm−3 in background and 13C-incubated cores) and decreased gradually with depth. Large differences were observed in the uptake of the algal material among species and between depth levels. During the experiment, 0.82 mg C m−2 were ingested, mainly by calcareous (~60%) and agglutinated (~40%) foraminifera. Uptake was highest at the sediment surface and 3 to 5 times less in deeper sediment horizons. Despite clear signs of vitality and a strong representation in the foraminiferal community, none of the soft-walled species showed a noticeable response to the offered algal material. We conclude that soft-walled foraminifera may not be important to the short-term phytodetrital matter cycling at the abyssal sea floor.