Ammonium excretion and oxygen respiration of tropical copepods and euphausiids exposed to oxygen minimum zone conditions

Diapycnal diffusivity in the core and oxycline of the tropical North Atlantic oxygen minimum zone

Abstract. A subsurface low oxygen zone is located in the eastern tropical North Atlantic Ocean (ETNA) in the upper ocean with the core of the hypoxic (O2 ≦60 µmol kg−1) oxygen minimum zone (OMZ) at 400 to 500 m depth. The subsurface circulation in the OMZ region is derived from observations and data assimilation results. Measurements in the ETNA of velocity, oxygen and of a tracer (CF3SF5) that was released in April 2008 at ∼ 8° N, 23° W (at ∼ 330 m depth) in November–December 2008, in November–December 2009 and October–November 2010 show the circulation in the upper part of the OMZ with spreading to the east in the North Equatorial Countercurrent (NECC) region and northwestward around the Guinea Dome. Three floats equipped with oxygen sensors deployed at ∼ 8° N, 23° W with parking depths at 330, 350 and 400 m depths were used to estimate velocity along the float trajectory at the surface and at the parking depth. At the 350 m park depth north of 9° N a cyclonic northwestward flow across the OMZ was observed. The northward drift of a float into the upper OMZ and a stronger cyclonic flow around the Guinea Dome seem to be connected to a strong Atlantic Meridional Mode (AMM) event in 2009. A near-surface cyclonic circulation cell east of the Cape Verde Islands reaches down into the OMZ layer. The circulation of the upper OMZ mirrors the near-surface circulation. Oxygen measurements from the cruises used here, as well as from other recent cruises up to the year 2014, confirm the continuous deoxygenation trend in the upper OMZ since the 1960s near the Guinea Dome. The three floats deployed with the tracer show spreading paths consistent with the overall observed tracer spreading. Oxygen sensors on the floats remained well calibrated for more than 20 months, and so the oxygen profiles can be used to investigate mesoscale eddy signatures. Mesoscale eddies may modify the oxygen distribution in OMZs. However, in general eddies are less energetic in the ETNA south of the Cape Verde Islands compared to similar latitudes in the eastern tropical South Pacific.

Distribution patterns of fragile gelatinous fauna in the open ocean remain scarcely documented. Using epi-and mesopelagic video transects in the eastern tropical North Atlantic, which features a mild but intensifying midwater oxygen minimum zone (OMZ), we established one of the first regional observations of diversity and abundance of large gelatinous zooplankton. We quantified the day and night vertical distribution of 46 taxa in relation to environmental conditions. While distribution may be driven by multiple factors, abundance peaks of individual taxa were observed in the OMZ core, both above and below the OMZ, only above, or only below the OMZ whereas some taxa did not have an obvious distribution pattern. In the eastern eropical North Atlantic, OMZ expansion in the course of global climate change may detrimentally impact taxa that avoid low oxygen concentrations (Beroe, doliolids), but favour taxa that occur in the OMZ (Lilyopsis, phaeodarians, Cydippida, Colobonema, Haliscera conica and Halitrephes) as their habitat volume might increase. While future efforts need to focus on physiology and taxonomy of pelagic fauna in the study region, our study presents biodiversity and distribution data for the regional epi- and mesopelagic zones of Cape Verde providing a regional baseline to monitor how climate change may impact the largest habitat on the planet, the deep pelagic realm.

Abstract. 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 ongoing 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) at 75 kHz (shipboard acoustic Doppler current profiler, ADCP) within the shallow OMZ of the eddy was evident compared to the nighttime distribution outside the eddy. Acoustic scatterers avoided the depth range between approximately 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, 300 kHz) 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 followed by zooplankton in response to in response to the eddy OMZ have been identified: (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.

Abstract. Ocean observations are analysed in the framework of Collaborative Research Center 754 (SFB 754) "Climate-Biogeochemistry Interactions in the Tropical Ocean" to study (1) the structure of tropical oxygen minimum zones (OMZs), (2) the processes that contribute to the oxygen budget, and (3) long-term changes in the oxygen distribution. The OMZ of the eastern tropical North Atlantic (ETNA), located between the well-ventilated subtropical gyre and the equatorial oxygen maximum, is composed of a deep OMZ at about 400 m in depth with its core region centred at about 20° W, 10° N and a shallow OMZ at about 100 m in depth, with the lowest oxygen concentrations in proximity to the coastal upwelling region off Mauritania and Senegal. The oxygen budget of the deep OMZ is given by oxygen consumption mainly balanced by the oxygen supply due to meridional eddy fluxes (about 60%) and vertical mixing (about 20%, locally up to 30%). Advection by zonal jets is crucial for the establishment of the equatorial oxygen maximum. In the latitude range of the deep OMZ, it dominates the oxygen supply in the upper 300 to 400 m and generates the intermediate oxygen maximum between deep and shallow OMZs. Water mass ages from transient tracers indicate substantially older water masses in the core of the deep OMZ (about 120–180 years) compared to regions north and south of it. The deoxygenation of the ETNA OMZ during recent decades suggests a substantial imbalance in the oxygen budget: about 10% of the oxygen consumption during that period was not balanced by ventilation. Long-term oxygen observations show variability on interannual, decadal and multidecadal timescales that can partly be attributed to circulation changes. In comparison to the ETNA OMZ, the eastern tropical South Pacific OMZ shows a similar structure, including an equatorial oxygen maximum driven by zonal advection but overall much lower oxygen concentrations approaching zero in extended regions. As the shape of the OMZs is set by ocean circulation, the widespread misrepresentation of the intermediate circulation in ocean circulation models substantially contributes to their oxygen bias, which might have significant impacts on predictions of future oxygen levels.

Vertical zonation and distributions of calanoid copepods through the lower oxycline of the Arabian Sea oxygen minimum zone

Abstract. Recent observations in the eastern tropical South Pacific (ETSP) have shown the key role of meso- and submesoscale processes (e.g. eddies) in shaping its hydrographic and biogeochemical properties. Off Peru, elevated primary production from coastal upwelling in combination with sluggish ventilation of subsurface waters fuels a prominent oxygen minimum zone (OMZ). Given that nitrous oxide (N2O) production–consumption processes in the water column are sensitive to oxygen (O2) concentrations, the ETSP is a region of particular interest to investigate its source–sink dynamics. To date, no detailed surveys linking mesoscale processes and N2O distributions as well as their relevance to nitrogen (N) cycling are available. In this study, we present the first measurements of N2O across three mesoscale eddies (two mode water or anticyclonic and one cyclonic) which were identified, tracked, and sampled during two surveys carried out in the ETSP in November–December 2012. A two-peak structure was observed for N2O, wherein the two maxima coincide with the upper and lower boundaries of the OMZ, indicating active nitrification and partial denitrification. This was further supported by the abundances of the key gene for nitrification, ammonium monooxygenase (amoA), and the gene marker for N2O production during denitrification, nitrite reductase (nirS). Conversely, we found strong N2O depletion in the core of the OMZ (O2 < 5 µmol L−1) to be consistent with nitrite (NO2−) accumulation and low levels of nitrate (NO3−), thus suggesting active denitrification. N2O depletion within the OMZ's core was substantially higher in the centre of mode water eddies, supporting the view that eddy activity enhances N-loss processes off Peru, in particular near the shelf break where nutrient-rich, productive waters from upwelling are trapped before being transported offshore. Analysis of eddies during their propagation towards the open ocean showed that, in general, “ageing” of mesoscale eddies tends to decrease N2O concentrations through the water column in response to the reduced supply of material to fuel N loss, although hydrographic variability might also significantly impact the pace of the production–consumption pathways for N2O. Our results evidence the relevance of mode water eddies for N2O distribution, thereby improving our understanding of the N-cycling processes, which are of crucial importance in times of climate change and ocean deoxygenation.

The genus Oithona has been considered the most abundant and ubiquitous copepod in the world's oceans. However, despite its importance, the metabolism of its developmental stages (nauplii and copepodites), crucial to explain their evolutionary success, is almost unknown. We determined respiration rates, ammonium and phosphate excretion rates, and the net growth efficiencies of early developmental stages of Oithona davisae as related to stage, body weight, temperature, and food availability. Respiration and excretion rates increased with increasing body weight and were positively related to temperature and food. Specific respiration rates of nauplii and copepodites varied from 0.11 to 0.55 d−1 depending on stage, body weight, temperature, and food availability. Metabolic C:N ratios were higher than 14, indicating lipid‐oriented metabolism. Assimilation efficiencies and net growth efficiencies ranged from 65% to 86% and from 23% to 32%, respectively, depending on body weight, stage, and temperature. Assimilation efficiencies and net growth efficiencies estimated using the respiration rates of nauplii with food were 1.7 times higher and 0.6 times lower, respectively, than those calculated using respiration rates of nauplii without food. Therefore, the use of respiration rates measured in filtered seawater led to substantial bias on the estimations of zooplankton carbon budget. O. davisae developmental stages exhibited similar assimilation and growth efficiencies but lower carbon‐specific respiratory losses than calanoid copepods. Hence, the low metabolic costs of Oithona compared with calanoids may be one reason for their success in marine ecosystems.

There is an incomplete description of the mid-depth circulation and its link to the oxygen minimum zone (OMZ) in the eastern tropical South Pacific. Subsurface currents of the OMZ in the eastern tropical South Pacific are investigated with a focus at 400 m depth, close to the core of the OMZ, using several Acoustic Doppler Current Profiler sections recorded in January and February 2009. Five profiling floats with oxygen sensors were deployed along 85°50’W in February 2009 with a drift depth at 400 m. Their spreading paths are compared with the model flow field from a 1/10° Tropical Pacific model (TROPAC01) and the Simple Ocean Data Assimilation (SODA) model. Overall the mean currents in the eastern tropical South Pacific are weak so that eddy variability influences the flow and ultimately feed oxygen-poor water to the OMZ. The center of the OMZ is a stagnant area so that floats stay much longer in this region and can even reverse direction. In one case of one float deployed at 8°S returned to the same location after 15 month. On the northern side of the OMZ in the equatorial current system, floats move rapidly to the west. Most current bands reported for the near surface layer exist also in the depth range of the OMZ. A schematic circulation flow field for the OMZ core depth is derived which shows the northern part of the South Pacific subtropical gyre south of the OMZ and the complicated zonal equatorial flow field north of the OMZ.

We measured the ammonium excretion, phosphate excretion and respiration rates of the scyphomedusa Aurelia aurita from Ondo Strait, in the central part of the Inland Sea of Japan, at 28 and 20°C. The rates measured at 28°C were converted to those at 20°C using the Q10 values, i.e. 1.56, 1.57 and 2.80, for ammonium excretion, phosphate excretion and respiration rates, respectively. The composite relationships between metabolic rates and wet weight of a medusa (WW, g, range 11–1330 g) at 20°C were expressed by the following allometric equations. For ammonium excretion rate (N, µmoles N medusa−1d−1): N = 0.497WW 1.09, phosphate excretion rate (P, µmoles P medusa−1d−1): P = 0.453WW 0.84, and respiration rate (R, µmoles O2 medusa−1d−1): R = 96.9WW 1.06. Mean O:N ratios (i.e. atomic ratios of 16.9 and 11.0 at 28 and 20°C, respectively) indicated that the metabolism of A. aurita medusae was protein-dominated. These metabolic parameters enabled us to estimate the nitrogen and phosphorus regeneration rates of an A. aurita medusa population typical of early summer in the Ondo Strait (means of water temperature, medusa individual weight and population biomass: 20°C, 200 g WW and 50.8 g WW m−3, respectively). Regenerated nitrogen and phosphorus were equivalent to 10.0 and 21.6% of phytoplankton uptake rates, respectively, nearly twice that estimated for mesozooplankton, demonstrating that A. aurita medusae are key components of the plankton community, influencing the trophic and nutrient dynamics in the Ondo Strait during early summer.

Herbivorous zooplankton species (Calanus plumchrus, Paracalanus parvus and Euphausia pacifica) and carnivorous species (Parathemisto pacifica and Pleurobrachia pileus) collected from Saanich Inlet, British Columbia, Canada, were maintained in the laboratory under fed and starved conditions. Respiration rate and excretion rates of ammonia and inorganic phosphate were measured successively on the same batch populations of each species in different feeding conditions. Respiration rate remained at a constant level or increased during the feeding experiment but decreased progressively in starved individuals. Herbivorous, but not carnivorous, species showed a rapid decrease in both excretion rates for the first few days of an experiment irrespective of feeding conditions. However, the general level of excretion rates of fed specimens was higher than that of starved ones. The O:N, N:P and O:P ratios were calculated from respiration, ammonia excretion and phosphate excretion and discussed in relation to metabolic substrates of animals during the experiment. A marked difference was shown in the O:N ratio between fed hervivores (>16) and fed carnivores (7 to 19), suggesting highly protein-oriented metabolism in the latter. One unknown factor causing variation in excretion rates is speculated to be the physiological stress on animals during sampling from the field. It is suggested that the laboratory measurement of realistic excretion rates of zooplankton is difficult owing to their large fluctuations, but this is not the case with respiration rate.

Abstract. Increasing deoxygenation (loss of oxygen) of the ocean, including expansion of oxygen minimum zones (OMZs), is a potentially important consequence of global warming. We examined present-day variability of vertical distributions of 23 calanoid copepod species in the Eastern Tropical North Pacific (ETNP) living in locations with different water column oxygen profiles and OMZ intensity (lowest oxygen concentration and its vertical extent in a profile). Copepods and hydrographic data were collected in vertically stratified day and night MOCNESS (Multiple Opening/Closing Net and Environmental Sensing System) tows (0–1000 m) during four cruises over a decade (2007–2017) that sampled four ETNP locations: Costa Rica Dome, Tehuantepec Bowl, and two oceanic sites further north (21–22∘ N) off Mexico. The sites had different vertical oxygen profiles: some with a shallow mixed layer, abrupt thermocline, and extensive very low oxygen OMZ core; and others with a more gradual vertical development of the OMZ (broad mixed layer and upper oxycline zone) and a less extensive OMZ core where oxygen was not as low. Calanoid copepod species (including examples from the genera Eucalanus, Pleuromamma, and Lucicutia) demonstrated different distributional strategies (implying different physiological characteristics) associated with this variability. We identified sets of species that (1) changed their vertical distributions and depth of maximum abundance associated with the depth and intensity of the OMZ and its oxycline inflection points; (2) shifted their depth of diapause; (3) adjusted their diel vertical migration, especially the nighttime upper depth; or (4) expanded or contracted their depth range within the mixed layer and upper part of the thermocline in association with the thickness of the aerobic epipelagic zone (habitat compression concept). These distribution depths changed by tens to hundreds of meters depending on the species, oxygen profile, and phenomenon. For example, at the lower oxycline, the depth of maximum abundance for Lucicutia hulsemannae shifted from ∼600 to ∼800 m, and the depth of diapause for Eucalanus inermis shifted from ∼500 to ∼775 m, in an expanded OMZ compared to a thinner OMZ, but remained at similar low oxygen levels in both situations. These species or life stages are examples of “hypoxiphilic” taxa. For the migrating copepod Pleuromamma abdominalis, its nighttime depth was shallow (∼20 m) when the aerobic mixed layer was thin and the low-oxygen OMZ broad, but it was much deeper (∼100 m) when the mixed layer and higher oxygen extended deeper; daytime depth in both situations was ∼300 m. Because temperature decreased with depth, these distributional depth shifts had metabolic implications. The upper ocean to mesopelagic depth range encompasses a complex interwoven ecosystem characterized by intricate relationships among its inhabitants and their environment. It is a critically important zone for oceanic biogeochemical and export processes and hosts key food web components for commercial fisheries. Among the zooplankton, there will likely be winners and losers with increasing ocean deoxygenation as species cope with environmental change. Changes in individual copepod species abundances, vertical distributions, and life history strategies may create potential perturbations to these intricate food webs and processes. Present-day variability provides a window into future scenarios and potential effects of deoxygenation.

Centennial to millennial-scale changes in oxygenation and productivity in the Eastern Tropical South Pacific during the last 25,000 years

Abstract. Repeat shipboard and multi-year moored observations obtained in the oxygen minimum zone (OMZ) of the eastern tropical North Atlantic (ETNA) were used to study the decadal change in oxygen for the period 2006–2015. Along 23° W between 6 and 14° N, oxygen decreased with a rate of −5.9 ± 3.5 µmol kg−1 decade−1 within the depth covering the deep oxycline (200–400 m), while below the OMZ core (400–1000 m) oxygen increased by 4.0 ± 1.6 µmol kg−1 decade−1 on average. The inclusion of these decadal oxygen trends in the recently estimated oxygen budget for the ETNA OMZ suggests a weakened ventilation of the upper 400 m, whereas the ventilation strengthened homogeneously below 400 m. The changed ventilation resulted in a shoaling of the ETNA OMZ of −0.03 ± 0.02 kg m−3 decade−1 in density space, which was only partly compensated by a deepening of isopycnal surfaces, thus pointing to a shoaling of the OMZ in depth space as well (−22 ± 17 m decade−1). Based on the improved oxygen budget, possible causes for the changed ventilation are analyzed and discussed. Largely ruling out other ventilation processes, the zonal advective oxygen supply stands out as the most probable budget term responsible for the decadal oxygen changes.

Mid-Holocene intensification of the oxygen minimum zone in the northeastern Arabian Sea