Description: Sampling was conducted during RV Meteor cruise M93 in austral summer 2013 in an area from 11ºS to 14ºS and approximately 120 km offshore to within 10 km of the Peruvian coast. Specimens were collected using a Hydrobios Multinet Maxi (0.5 m2 mouth opening, 330 µm mesh size, 9 nets) and a WP-2 net (Hydrobios, 0.26 m2 mouth opening, 200 µm mesh size). P. monodon were identified according to http://researchdata.museum.vic.gov.au/squatlobster/delta/deltakey.html. Specimens were transferred into filtered, well-oxygenated seawater immediately after the catch and maintained for 4 to 16 hours prior to physiological experiments. Maintenance and physiological experiments were conducted at 13°C as the temperature observed at 100 to 200 m depth in the OMZ ranged from 13.7 to 12.7°C.

Description: Respiration and ammonium excretion rates at different oxygen partial pressure were measured for calanoid copepods and euphausiids from the Eastern Tropical South Pacific and the Eastern Tropical North Atlantic. All specimens used for experiments were caught in the upper 400 m of the water column and only animals appearing unharmed and fit were used for experiments. Specimens were sorted, identified and transferred into aquaria with filtered, well-oxygenated seawater immediately after the catch and maintained for 1 to 13 hours prior to physiological experiments at the respective experimental temperature. Maintenance and physiological experiments were conducted in darkness in temperature-controlled incubators at 11, 13 or 23 degree C (±1). Before and during experiments, animals were not fed. Respiration and ammonium excretion rate measurements (both in µmol h-1 gDW-1) at varying oxygen concentrations were conducted in 12 to 60 mL gas-tight glass bottles. These were equipped with oxygen microsensors (ø 3 mm, PreSens Precision Sensing GmbH, Regensburg, Germany) attached to the inner wall of the bottles to monitor oxygen concentrations non-invasively. Read-out of oxygen concentrations was conducted using multi-channel fiber optic oxygen transmitters (Oxy-4 and Oxy-10 mini, PreSens Precision Sensing GmbH, Regensburg, Germany) that were connected via optical fibers to the outside of the bottles directly above the oxygen microsensor spots. Measurements were started at pre-adjusted oxygen and carbon dioxide levels. For this, seawater stocks with adjusted pO2 and pCO2 were prepared by equilibrating 3 to 4 L of filtered (0.2 µm filter Whatman GFF filter) and UV - sterilized (Aqua Cristal UV C 5 Watt, JBL GmbH & Co. KG, Neuhofen, Germany) water with premixed gases (certified gas mixtures from Air Liquide) for 4 hours at the respective experimental temperature. pCO2 levels were chosen to mimic the environmental pCO2 in the ETSP OMZ or the ETNA OMZ. Experimental runs were conducted with 11 to 15 trial incubations (1 or 2 animals per incubation bottle and three different treatment levels) and three animal-free control incubations (one per experimental treatment). During each run, experimental treatments comprised 100% air saturation as well as one reduced air saturation level with and without CO2. Oxygen concentrations in the incubation bottles were recorded every 5 min using the fiber-optic microsensor system and data recording for respiration rate determination was started immediately after all animals were transferred. Respiration rates were calculated from the slope of oxygen decrease over selected time intervals. Chosen time intervals were 20 to 105 min long. No respiration rate was calculated for the first 20 to 60 min after animal transfer to avoid the impact of enhanced activity of the animal or changes in the bottle water temperature during initial handling on the respiration rates and oxygen readings. Respiration rates were obtained over a maximum of 16 hours incubation time and slopes were linear at normoxia to mild hypoxia. Respiration rates in animal-free control bottles were used to correct for microbial activity. These rates were 〈 2% of animal respiration rates at normoxia. Samples for the measurement of ammonium concentrations were taken after 2 to 10 hours incubation time. Ammonium concentration was determined fluorimetrically (Holmes et al., 1999). Ammonium excretion was calculated as the concentration difference between incubation and animal-free control bottles. Some specimens died during the respiration and excretion rate measurements, as indicated by a cessation of respiration. No excretion rate measurements were conducted in this case, but the oxygen level at which the animal died was noted.

Description: High primary productivity in the equatorial Atlantic and Pacific oceans is one of the key features of tropical ocean biogeochemistry and fuels a substantial flux of particulate matter towards the abyssal ocean. How biological processes and equatorial current dynamics shape the particle size distribution and flux, however, is poorly understood. Here we use high-resolution size-resolved particle imaging and Acoustic Doppler Current Profiler data to assess these influences in equatorial oceans. We find an increase in particle abundance and flux at depths of 300 to 600 m at the Atlantic and Pacific equator, a depth range to which zooplankton and nekton migrate vertically in a daily cycle. We attribute this particle maximum to faecal pellet production by these organisms. At depths of 1,000 to 4,000 m, we find that the particulate organic carbon flux is up to three times greater in the equatorial belt (1° S–1° N) than in off-equatorial regions. At 3,000 m, the flux is dominated by small particles less than 0.53 mm in diameter. The dominance of small particles seems to be caused by enhanced active and passive particle export in this region, as well as by the focusing of particles by deep eastward jets found at 2° N and 2° S. We thus suggest that zooplankton movements and ocean currents modulate the transfer of particulate carbon from the surface to the deep ocean.

Description: We estimated the relative contribution of atmospheric Nitrogen (N) input (wet and dry deposition and N fixation) to the epipelagic food web by measuring N isotopes of different functional groups of epipelagic zooplankton along 23°W (17°N-4°S) and 18°N (20-24°W) in the Eastern Tropical Atlantic. Results were related to water column observations of nutrient distribution and vertical diffusive flux as well as colony abundance of Trichodesmium obtained with an Underwater Vision Profiler (UVP5). The thickness and depth of the nitracline and phosphocline proved to be significant predictors of zooplankton stable N isotope values. Atmospheric N input was highest (61% of total N) in the strongly stratified and oligotrophic region between 3 and 7°N, which featured very high depth-integrated Trichodesmium abundance (up to 9.4×104 colonies m-2), strong thermohaline stratification and low zooplankton delta15N (~2 per mil). Relative atmospheric N input was lowest south of the equatorial upwelling between 3 and 5°S (27%). Values in the Guinea Dome region and north of Cape Verde ranged between 45 and 50%, respectively. The microstructure-derived estimate of the vertical diffusive N flux in the equatorial region was about one order of magnitude higher than in any other area (approximately 8 mmol m-2 d 1). At the same time, this region received considerable atmospheric N input (35% of total). In general, zooplankton delta15N and Trichodesmium abundance were closely correlated, indicating that N fixation is the major source of atmospheric N input. Although Trichodesmium is not the only N fixing organism, its abundance can be used with high confidence to estimate the relative atmospheric N input in the tropical Atlantic (r2 = 0.95). Estimates of absolute N fixation rates are two- to tenfold higher than incubation-derived rates reported for the same regions. Our approach integrates over large spatial and temporal scales and also quantifies fixed N released as dissolved inorganic and organic N. In a global analysis, it may thus help to close the gap in oceanic N budgets.

Description: The eastern tropical North Atlantic (ETNA) features a mesopelagic oxygen minimum zone (OMZ) at approximately 300-600 m depth. Here, oxygen concentrations rarely fall below 40 µmol O2 kg-1, but are expected to decline under future projections of global warming. The recent discovery of mesoscale eddies that harbour a shallow suboxic (〈5 µmol O2 kg-1) OMZ just below the mixed layer could serve to identify zooplankton groups that may be negatively or positively affected by on-going ocean deoxygenation. In spring 2014, a detailed survey of a suboxic anticyclonic modewater eddy (ACME) was carried out near the Cape Verde Ocean Observatory (CVOO), combining acoustic and optical profiling methods with stratified multinet hauls and hydrography. The multinet data revealed that the eddy was characterized by an approximately 1.5-fold increase in total area-integrated zooplankton abundance. At nighttime, when a large proportion of acoustic scatterers is ascending into the upper 150 m, a drastic reduction in mean volume backscattering (Sv, shipboard ADCP, 75kHz) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers were avoiding the depth range between about 85 to 120 m, where oxygen concentrations were lower than approximately 20 µmol O2 kg-1, indicating habitat compression to the oxygenated surface layer. This observation is confirmed by time-series observations of a moored ADCP (upward looking, 300kHz) during an ACME transit at the CVOO mooring in 2010. Nevertheless, part of the diurnal vertical migration (DVM) from the surface layer to the mesopelagic continued through the shallow OMZ. Based upon vertically stratified multinet hauls, Underwater Vision Profiler (UVP5) and ADCP data, four strategies have been identified to be followed by zooplankton in response to the eddy OMZ: i) shallow OMZ avoidance and compression at the surface (e.g. most calanoid copepods, euphausiids), ii) migration to the shallow OMZ core during daytime, but paying O2 debt at the surface at nighttime (e.g. siphonophores, Oncaea spp., eucalanoid copepods), iii) residing in the shallow OMZ day and night (e.g. ostracods, polychaetes), and iv) DVM through the shallow OMZ from deeper oxygenated depths to the surface and back. For strategy i), ii) and iv), compression of the habitable volume in the surface may increase prey-predator encounter rates, rendering zooplankton and micronekton more vulnerable to predation and potentially making the eddy surface a foraging hotspot for higher trophic levels. With respect to long-term effects of ocean deoxygenation, we expect avoidance of the mesopelagic OMZ to set in if oxygen levels decline below approximately 20 µmol O2 kg-1. This may result in a positive feedback on the OMZ oxygen consumption rates, since zooplankton and micronekton respiration within the OMZ as well as active flux of dissolved and particulate organic matter into the OMZ will decline.

Description: In ocean waters, anaerobic microbial respiration should be confined to the anoxic waters found in coastal regions and tropical oxygen minimum zones, where it is energetically favourable. However, recent molecular and geochemical evidence has pointed to a much broader distribution of denitrifying and sulfate-reducing microbes. Anaerobic metabolisms are thought to thrive in microenvironments that develop inside sinking organic aggregates, but the global distribution and geochemical significance of these microenvironments is poorly understood. Here, we develop a new size-resolved particle model to predict anaerobic respiration from aggregate properties and seawater chemistry. Constrained by observations of the size spectrum of sinking particles, the model predicts that denitrification and sulfate reduction can be sustained throughout vast, hypoxic expanses of the ocean, and could explain the trace metal enrichment observed in particles due to sulfide precipitation. Globally, the expansion of the anaerobic niche due to particle microenvironments doubles the rate of water column denitrification compared with estimates based on anoxic zones alone, and changes the sensitivity of the marine nitrogen cycle to deoxygenation in a warming climate.

Description: Upwelling systems play a key role in the global carbon and nitrogen cycles and are also of local relevance due to their high productivity and fish resources. To capture and understand the high spatial and temporal variability of physical and biogeochemical parameters found in these regions novel measurement technics have to be combined in an interdisciplinary manner. Here we use high-resolution glider-based physical-biogeochemical observations in combination with ship-based underwater vision profiler, sensor and bottle data to investigate the drivers of oxygen and nitrate variability across the shelf break off Mauritania in June 2014. Distinct oxygen and nitrate variability shows up in our glider data. High oxygen and low nitrate anomalies were clearly related to water mass variability and probably linked to ocean transport. Low oxygen and high nitrate patches co-occurred with enhanced turbidity signals close to the seabed, which suggests locally high microbial respiration of resuspended organic matter near the sea floor. This interpretation is supported by high particle abundance observed by the underwater vision profiler and enhanced particle-based respiration rate estimates close to the seabed. Discrete in-situ measurements of dissolved organic carbon and amino acids suggest the formation of dissolved organic carbon due to particle dissolution near the seabed fueling additional microbial respiration. Our high-resolution interdisciplinary observations highlight the complex interplay of remote and local physical-biogeochemical drivers of oxygen and nitrate variability off Mauritania, which cannot be captured by classical shipboard observations alone.

Description: Gelatinous zooplankton hold key functions in the ocean and have been shown to significantly influence the transport of organic carbon to the deep sea. We discovered a gelatinous, flux‐feeding polychaete of the genus Poeobius in very high abundances in a mesoscale eddy in the tropical Atlantic Ocean, where it co‐occurred with extremely low particle concentrations. Subsequent analysis of an extensive in situ imaging dataset revealed that Poeobius sp. occurred sporadically between 5°S–20°N and 16°W–46°W in the upper 1000 m. Abundances were significantly elevated and the depth distribution compressed in anticyclonic modewater eddies (ACMEs). In two ACMEs, high Poeobius sp. abundances were associated with strongly reduced particle concentrations and fluxes in the layers directly below the polychaete. We discuss possible reasons for the elevated abundances of Poeobius sp. in ACMEs and provide estimations showing that a single zooplankton species can completely intercept the downward particle flux by feeding with their mucous nets, thereby substantially altering the biogeochemical setting within the eddy.

Description: A Hydrobios Multinet Midi with an aperture of 0.25 m2 and 5 nets (mesh size 200 μm) was deployed for vertically stratified hauls on several cruises, mostly in paired day-night hauls to quantify diel vertical migration. Standard depths used for these deployments were 1000-600-300-200-100-0 m. On M93, a Multinet Maxi (9 nets, 333 μm mesh) was used instead. Samples were fixated in 4% formaldehyde in seawater solution, scanned at GEOMAR or at OSCM, and analyzed using automated imaging software (Gorsky et al., 2010) allowing taxonomical classification as well as the estimation of taxon-specific biomass (Lehette and Hernandez-Leon, 2006) and metabolic rates.