Eastern Tropical South Pacific Ocean Research Articles

The 1991–2020 sea surface temperature normals

AbstractThe 1991–2020 climate normals for sea surface temperature (SST) are computed based on the NOAA Daily Optimum Interpolation SST dataset. This is the first time that high‐resolution SST normals with global coverage can be achieved in the satellite SST era. Normals are one of the fundamental parameters in describing and understanding weather and climate and provide decision‐making information to industry, public, and scientific communities. This product suite includes SST mean, standard deviation, count and extreme parameters at daily, monthly, seasonal and annual time scales on 0.25° spatial grids. The main feature of the SST mean state revealed by the normals is that in the Tropics, the Indo‐Pacific Ocean is dominated by the warm pool (SST ≥ 28°C) while the eastern Pacific is characterized by the cold tongue (SST ≤ 24°C); in the midlatitudes, SSTs are in zonal patterns with high meridional gradients. Daily SST standard deviations are generally small (<1.0°C) except in frontal zones (>1.5°C) mostly associated with ocean currents such as the Gulf Stream, Kuroshio and Equatorial Currents. Compared to the 1982–2011 climatology, the 1991–2020 mean SSTs increased over most global areas but obvious cooling is seen in the Southern Ocean, eastern tropical South Pacific Ocean and North Atlantic warming hole. The Indo‐Pacific warm pool (IPWP) is found to have strengthened in both intensity and coverage since 1982–2011. By a count parameter criterion of ≥300 days annually with SST ≥ 28°C, the IPWP coverage increased 33% from 1982–2011 to 1991–2020. The global mean SST of 1991–2020 is warmer than that of 1982–2011, and the warming rate over 1991–2020 doubles that over 1901–2020.

Marine N2O cycling from high spatial resolution concentration, stable isotopic and isotopomer measurements along a meridional transect in the eastern Pacific Ocean

Nitrous oxide (N2O) is a potent greenhouse gas and ozone depleting substance, with the ocean accounting for about one third of global emissions. In marine environments, a significant amount of N2O is produced by biological processes in Oxygen Deficient Zones (ODZs). While recent technological advances are making surface N2O concentration more available, high temporal and spatial resolution water-column N2O concentration data are relatively scarce, limiting global N2O ocean models’ predictive capability. We present a N2O concentration, stable isotopic composition and isotopomer dataset of unprecedently large spatial coverage and depth resolution in the broader Pacific, crossing both the eastern tropical South and North Pacific Ocean ODZs collected as part of the GO-SHIP P18 repeat hydrography program in 2016/2017. We complement these data with dissolved gases (nitrogen, oxygen, argon) and nitrate isotope data to investigate the pathways controlling N2O production in relation to apparent oxygen utilization and fixed nitrogen loss. N2O yield significantly increased under low oxygen conditions near the ODZs. Keeling plot analysis revealed different N2O sources above the ODZs under different oxygen regimes. Our stable isotopic data and relationships between the N2O added by microbial processes (ΔN2O) and dissolved inorganic nitrogen (DIN) deficit confirm increased N2O production by denitrification under low oxygen conditions near the oxycline where the largest N2O accumulations were observed. The slope for δ18O-N2O versus site preference (SP, the difference between the central (α) and outer (β) N atoms in the linear N2O molecule) in the eastern tropical North Pacific ODZ was lower than expected for pure N2O reduction, likely because of the observed decrease in δ15Nβ. This trend is consistent with prior ODZ studies and attributed to concurrent production of N2O from nitrite with a low δ15N or denitrification with a SP >0‰. We estimated apparent isotope effects for N2O consumption in the ETNP ODZ of 3.6‰ for 15Nbulk, 9.4‰ for 15Nα, -2.3‰ for 15Nβ, 12.0‰ for 18O, and 11.7‰ for SP. These values were generally within ranges previously reported for previous laboratory and field experiments.

Diving deeper: Mesopelagic fish biomass estimates comparison using two different models

A growing population on a planet with limited resources demands finding new sources of protein. Hence, fisheries are turning their perspectives towards mesopelagic fish, which have, so far, remained relatively unexploited and poorly studied. Large uncertainties are associated with regards to their biomass, turn-over rates, susceptibility to environmental forcing and ecological and biogeochemical role. Models are useful to disentangle sources of uncertainties and to understand the impact of different processes on the biomass. In this study, we employed two food-web models – OSMOSE and the model by Anderson et al. (2019, or A2019) – coupled to a regional physical–biogeochemical model to simulate mesopelagic fish in the Eastern Tropical South Pacific ocean. The model by A2019 produced the largest biomass estimate, 26 to 130% higher than OSMOSE depending on the mortality parameters used. However, OSMOSE was calibrated to match observations in the coastal region off Peru and its temporal variability is affected by an explicit life cycle and food web. In contrast, the model by A2019 is more convenient to perform uncertainty analysis and it can be easily coupled to a biogeochemical model to estimate mesopelagic fish biomass. However, it is based on a flow analysis that had been previously applied to estimate global biomass of mesopelagic fish but has never been calibrated for the Eastern Tropical South Pacific. Furthermore, it assumes a steady-state in the energy transfer between primary production and mesopelagic fish, which may be an oversimplification for this highly dynamic system. OSMOSE is convenient to understand the interactions of the ecosystem and how including different life stages affects the model response. The combined strengths of both models allow us to study mesopelagic fish from a holistic perspective, taking into account energy fluxes and biomass uncertainties based on primary production, as well as complex ecological interactions.

Moisture Source for the Precipitation of Tropical Cyclones over the Pacific Ocean through a Lagrangian Approach

Abstract Tropical cyclones (TCs) are an important component of the hydrological cycle at tropical latitudes. In this study, we investigated the origin of precipitation associated with TCs formed from 1980 to 2018 over the Pacific Ocean in three subbasins: the western North Pacific Ocean (WNP), central and east Pacific Ocean (NEPAC), and South Pacific Ocean (SPO) basins. The analysis was performed throughout the TC lifetime during genesis, when they reached the lifetime maximum intensity (LMI), and the dissipation stage. The backward trajectories of all precipitant atmospheric parcels residing over the TC locations from the global outputs of the Lagrangian Flexible Particle (FLEXPART) dispersion model fed by the ERA-Interim dataset were used to identify moisture sources. The South and East China Seas and the western tropical North Pacific Ocean were identified as the principal moisture sources in the WNP basin, while the atmospheric moisture that precipitated mainly came from the eastern tropical North and South Pacific Ocean in the NEPAC basin, followed by the Caribbean Sea. Meanwhile, the Coral Sea, western tropical South Pacific Ocean, and northern Australia are the origins of the moisture in the SPO. The mean moisture uptake per TC was higher during the hurricane category than during any other stage in each basin.

The cyanobacterium Prochlorococcus has divergent light-harvesting antennae and may have evolved in a low-oxygen ocean

Marine picocyanobacteria of the genus Prochlorococcus are the most abundant photosynthetic organisms in the modern ocean, where they exert a profound influence on elemental cycling and energy flow. The use of transmembrane chlorophyll complexes instead of phycobilisomes as light-harvesting antennae is considered a defining attribute of Prochlorococcus Its ecology and evolution are understood in terms of light, temperature, and nutrients. Here, we report single-cell genomic information on previously uncharacterized phylogenetic lineages of this genus from nutrient-rich anoxic waters of the eastern tropical North and South Pacific Ocean. The most basal lineages exhibit optical and genotypic properties of phycobilisome-containing cyanobacteria, indicating that the characteristic light-harvesting antenna of the group is not an ancestral attribute. Additionally, we found that all the indigenous lineages analyzed encode genes for pigment biosynthesis under oxygen-limited conditions, a trait shared with other freshwater and coastal marine cyanobacteria. Our findings thus suggest that Prochlorococcus diverged from other cyanobacteria under low-oxygen conditions before transitioning from phycobilisomes to transmembrane chlorophyll complexes and may have contributed to the oxidation of the ancient ocean.

Timescales of ventilation and consumption of oxygen and fixed nitrogen in the eastern tropical South Pacific oxygen deficient zone from transient tracers

The anthropogenic trace gases chlorofluorocarbon (CFC)-12 and sulfur hexafluoride (SF6) were measured during 2013 in the eastern tropical South Pacific Ocean (ETSP) offshore Chile and Peru (12°-22°S, 70°-86°W). Since the WOCE P21 line along ~17°S in 1993, the CFC-12 penetration depth increased from ~550 m to ~800 m. In 2013, CFC-12 had penetrated through the bottom of the oxygen deficient zone (ODZ, where oxygen (O2) < 4.5 μmol kg−1) at all stations, indicating that a portion of waters in this ODZ are ventilated on timescales < 60 years. Isopycnal trends in pSF6 and pCFC-12 ages versus AOU indicated oxygen utilization rates of 11.2 ± 4.7 μmol kg−1 yr−1 just above the ODZ (90–130 m) and 1.0 ± 0.5 μmol kg−1 yr−1 beneath the ODZ (400–700 m). Isopycnal trends in pSF6 ages and nutrients implied fixed N-loss rates of 0.6 ± 0.4 μmol kg−1 yr−1 at the top of the ODZ (~120 m). The pSF6 and pCFC-12 ages were significantly younger than mean ages estimated from one-dimensional transit time distributions, which were difficult to constrain using the SF6 and CFC-12 tracer combination. Despite the fact that tracer concentrations tend to underestimate mean ages, and thus overestimate nutrient regeneration/consumption rates, N-loss rates were undetectable (<0.5 μmol kg−1 yr−1) within the ODZ itself (~175–400 m). When integrated over depth, the oxygen and nitrogen consumption rates determined above and below the ODZ implied total organic carbon (C) remineralization rates on the order of 0.6 ± 0.1 mol C m−2 yr−1. These low C-export rates, and the decadal ventilation timescale of this ODZ, support a body of work suggesting that the ODZ may be sustained by inputs of high-tracer, low-oxygen waters from the adjacent Peru-Chile coastal upwelling system rather than by organic matter oxidation occurring locally.

Estimating fixed nitrogen loss and associated isotope effects using concentration and isotope measurements of NO3–, NO2–, and N2 from the Eastern Tropical South Pacific oxygen deficient zone

Quantifying the pathways of fixed nitrogen (N) loss in marine oxygen deficient zones (ODZs) and the isotopic fractionation caused by these processes are important for understanding the marine fixed N budget and its potential for change. In this study, a variety of approaches were used to quantify fixed N loss in the eastern tropical South Pacific Ocean (ETSP). The required measurements included nutrient concentration (nitrate—NO3-, nitrite—NO2-, and phosphate—PO43-), gas ratio (N2/Ar) measurements, and stable N and O isotopes in NO3-, NO2-, and nitrogen gas (N2). The dissolved inorganic nitrogen deficit calculated from [PO43-] ([DIN]def,P) exceeded the concentration of N2 gas biologically produced in the ODZ (local [N2]bio) throughout the ODZ at most stations, likely due to release of PO43- from sediments driving up [DIN]def,P. Calculating DIN deficit using water mass analysis and local oxygen (O2) consumption ([DIN]def,OMP) yielded better agreement with local [N2]bio than [DIN]def,P, except at the maximum [N2]bio, where [DIN]def,OMP misses contributions of anaerobic ammonia oxidation (anammox) to N2 production. We used the mismatch between [DIN]def,OMP and [N2]bio to estimate a 29% contribution of anammox to [N2]bio. Stable isotopic measurements of NO2-, NO3-, and N2 were used alongside [N2]bio and new estimates of [DIN]def to calculate N and O isotope effects for NO3- reduction (15εNAR and 18εNAR, respectively), and N isotope effects for DIN removal (15εDIN-R). While the various methods for estimating [DIN]def had little effect on the isotope effects for DIN removal, differences between 15εNAR and 15εDIN-R, and variations with depth in the ODZ were observed. Using a simple time-dependent ODZ model, we interpreted these patterns to reflect the influences of NO2- oxidation and NO2- accumulation on expression of isotopic fractionation in the ODZ.

Nitric oxide (NO) in the oxygen minimum zone off Peru

Nitric oxide (NO) is a short-lived compound of the marine nitrogen cycle. However, measurements of NO in seawater are analytically challenging and our knowledge about its oceanic distribution is, therefore, rudimentary. NO was measured in the oxygen minimum zone (OMZ) of the eastern tropical South Pacific Ocean (ETSP) off Peru during R/V Meteor cruise M93 in February/March 2013. NO concentrations ranged from close to or below the detection limit (0.5 nmol L−1) in the surface layer to 9.5 nmol L−1 in the OMZ. NO concentrations increased significantly when oxygen (O2) concentrations dropped below 1–2 µmol L−1. We found positive correlations between NO and NO2- as well as between NO and the abundance of archaeal amoA, a marker gene for archaeal nitrifiers. No trends between NO and nirS and hzo, marker genes for canonical denitrification and anammox, respectively, were found. To this end, we conclude that NO off Peru was mainly produced by archaeal nitrifier-denitrification at low O2 concentrations in the OMZ.

Radium-228 as a tracer of dissolved trace element inputs from the Peruvian continental margin

Abstract Continental margins play a central role in the composition of seawater by being an important source of trace element essentials to the functioning of the ocean ecosystems. Here, we measured long-lived radium isotopes ( 226 Ra, 228 Ra) along a zonal transect at 12°S (US GEOTRACES GP16) in the eastern tropical South Pacific Ocean. We used 228 Ra to quantify the trace element and isotope (TEI) fluxes (DMn, DFe, and DCo) delivered from the Peruvian continental i) shelf and ii) slope. First, elevated 228 Ra activities were measured in surface water over the entire transect (~ 8500 km), evidence that the continental shelf is an important source of sediment-derived TEIs not only to coastal areas, but to central Pacific Ocean waters. Modeled 228 Ra shelf fluxes combined with water column dissolved TEI/ 228 Ra ratios were used to quantify the shelf-ocean input rates (normalized to shelf-area) for DMn (3.3 × 10 3 μmol m − 2 y − 1 ), DFe (1.5 × 10 3 μmol m − 2 y − 1 ), and DCo (1.0 × 10 2 μmol m − 2 y − 1 ). Second, co-occurring plumes of 228 Ra, DFe, and DMn extended over 1800 km from the margin at 1000–2500 m depth, indicative of a continental slope sediment TEI input to the intermediate water column. The 228 Ra gradient allowed us to derive an effective horizontal eddy diffusion coefficient (K h ) of 46 m 2 s − 1 , which in turn permitted the calculation of slope sediment DMn (6.4 μmol m − 2 y − 1 ) and DFe (5.9 × 10 2 μmol m − 2 y − 1 ) fluxes based on their offshore concentration gradients. On the scale of the South Pacific continental margin between 0–20°S, the DMn shelf flux is approximately 2–3 orders of magnitude higher than the slope flux, while the DFe shelf/slope flux is ~ 3:1. Both shelf and slope sediment derived DMn was transported over a significant distance towards the ocean interior, while DFe concentration gradients were steep, consistent with longer water column residence time for DMn as compared to DFe in marine systems. These findings highlight the importance of considering the continental slope-ocean boundary in the oceanic budgets of biologically-important trace elements.

Linking diverse nutrient patterns to different water masses within anticyclonic eddies in the upwelling system off Peru

Abstract. Ocean eddies can both trigger mixing (during their formation and decay) and effectively shield water encompassed from being exchanged with ambient water (throughout their lifetimes). These antagonistic effects of eddies complicate the interpretation of synoptic snapshots typically obtained by ship-based oceanographic measurement campaigns. Here we use a coupled physical–biogeochemical model to explore biogeochemical dynamics within anticyclonic eddies in the eastern tropical South Pacific Ocean. The goal is to understand the diverse biogeochemical patterns that have been observed at the subsurface layers of the anticyclonic eddies in this region. Our model results suggest that the diverse subsurface nutrient patterns within eddies are associated with the presence of water masses of different origins at different depths.

Distribution of mercury species across a zonal section of the eastern tropical South Pacific Ocean (U.S. GEOTRACES GP16)

Monomethylmercury is a neurotoxin formed in the ocean that accumulates in fish. To better understand the chemical cycling of mercury (Hg) in the ocean, multiple Hg species were measured during the 2013 U.S. GEOTRACES section in the eastern tropical South Pacific Ocean (GP16) from Peru to Tahiti. Total Hg (HgT), elemental Hg (Hg0), monomethyl-Hg (MMHg), and dimethyl-Hg (DMHg) were measured in high resolution across the section, which included productive surface waters in the Peru upwelling region, a suboxic oxygen minimum zone (OMZ), and an expansive hydrothermal vent plume from the East Pacific Rise (EPR). Filtered HgT accumulated in upwelled waters near the coast of Peru where oxygen concentrations were the lowest, and MMHg represented 10–20% of the total Hg upwelling flux. The buoyant hydrothermal plume was not enriched with Hg; however, water below 2500m had anomalously high concentrations of filtered HgT beneath the plume and in warmer bottom waters in the eastern part of the section. Elemental Hg averaged <10% of HgT in the water column and concentrations were greatest near the Peru margin. Concentrations of MMHg and DMHg were greatest in low-oxygen thermocline waters, and in the upwelling region, vertical maxima of MMHg were found at the top, core, and bottom of the suboxic OMZ. The MMHg:DMHg molar ratio was variable throughout the upper water column, however, DMHg was the dominant form of methylated Hg in deep water masses. This section is the first examination of Hg speciation and distributions near the Peru coast, a region that supports a highly productive fishery and a potential source of MMHg to consumers of seafood.

N&amp;lt;sub&amp;gt;2&amp;lt;/sub&amp;gt; fixation in eddies of the eastern tropical South Pacific Ocean

Abstract. Mesoscale eddies play a major role in controlling ocean biogeochemistry. By impacting nutrient availability and water column ventilation, they are of critical importance for oceanic primary production. In the eastern tropical South Pacific Ocean off Peru, where a large and persistent oxygen-deficient zone is present, mesoscale processes have been reported to occur frequently. However, investigations into their biological activity are mostly based on model simulations, and direct measurements of carbon and dinitrogen (N2) fixation are scarce.We examined an open-ocean cyclonic eddy and two anticyclonic mode water eddies: a coastal one and an open-ocean one in the waters off Peru along a section at 16° S in austral summer 2012. Molecular data and bioassay incubations point towards a difference between the active diazotrophic communities present in the cyclonic eddy and the anticyclonic mode water eddies.In the cyclonic eddy, highest rates of N2 fixation were measured in surface waters but no N2 fixation signal was detected at intermediate water depths. In contrast, both anticyclonic mode water eddies showed pronounced maxima in N2 fixation below the euphotic zone as evidenced by rate measurements and geochemical data. N2 fixation and carbon (C) fixation were higher in the young coastal mode water eddy compared to the older offshore mode water eddy. A co-occurrence between N2 fixation and biogenic N2, an indicator for N loss, indicated a link between N loss and N2 fixation in the mode water eddies, which was not observed for the cyclonic eddy. The comparison of two consecutive surveys of the coastal mode water eddy in November 2012 and December 2012 also revealed a reduction in N2 and C fixation at intermediate depths along with a reduction in chlorophyll by half, mirroring an aging effect in this eddy. Our data indicate an important role for anticyclonic mode water eddies in stimulating N2 fixation and thus supplying N offshore.

Low rates of nitrogen fixation in eastern tropical South Pacific surface waters

An extensive region of the Eastern Tropical South Pacific (ETSP) Ocean has surface waters that are nitrate-poor yet phosphate-rich. It has been proposed that this distribution of surface nutrients provides a geochemical niche favorable for N2fixation, the primary source of nitrogen to the ocean. Here, we present results from two cruises to the ETSP where rates of N2fixation and its contribution to export production were determined with a suite of geochemical and biological measurements. N2fixation was only detectable using nitrogen isotopic mass balances at two of six stations, and rates ranged from 0 to 23 µmol N m(-2)d(-1)based on sediment trap fluxes. Whereas the fractional importance of N2fixation did not change, the N2-fixation rates at these two stations were several-fold higher when scaled to other productivity metrics. Regardless of the choice of productivity metric these N2-fixation rates are low compared with other oligotrophic locations, and the nitrogen isotope budgets indicate that N2fixation supports no more than 20% of export production regionally. Although euphotic zone-integrated short-term N2-fixation rates were higher, up to 100 µmol N m(-2)d(-1), and detected N2fixation at all six stations, studies of nitrogenase gene abundance and expression from the same cruises align with the geochemical data and together indicate that N2fixation is a minor source of new nitrogen to surface waters of the ETSP. This finding is consistent with the hypothesis that, despite a relative abundance of phosphate, iron may limit N2fixation in the ETSP.

Diazotroph community structure in the deep oxygen minimum zone of the Costa Rica Dome.

Oxygen minimum zones (OMZs), characterized by depleted dissolved oxygen concentration in the intermediate depth of the water column, are predicted to expand under the influence of global warming. Recent studies in the Eastern Tropical South Pacific Ocean and Arabian Sea have reported that heterotrophic nitrogen fixation is active in the OMZs. In this study, we investigated the community structure of diazotrophs in the OMZ of the Costa Rica Dome (CRD) upwelling region in the Eastern Tropical North Pacific Ocean, using 454-pyrosequencing of nifH gene amplicons. Comparing diazotroph assemblages in different depth strata of the OMZ (200-1000 m in depth), we found a unique diazotroph community in the OMZ core, which was mainly dominated by methanotroph-like diazotrophs, suggesting a potential coupling of nitrogen cycle and methane assimilation. In addition, some OTUs revealed in this study, especially those belonging to the large sub-cluster Vibrio diazotrophicus, were reported to be abundant and expressing the nifH gene in other OMZs. Our results suggest that the unique hydrographic conditions in OMZs may support similar assemblages of diazotrophs, and heterotrophic nitrogen fixation could also be occurring in our studied region. Our study provides the first insight into the composition and distribution of putative diazotrophs in the CRD OMZ.

An improved method for the determination of dissolved nitric oxide (NO) in seawater samples

Abstract. Nitric oxide (NO) is a short-lived intermediate of the oceanic nitrogen cycle; however, due to its high reactivity, measurements of dissolved NO in seawater are rare. Here we present an improved method to determine NO concentrations in discrete seawater samples. The set-up of our system consisted of a chemiluminescence NO analyser connected to a stripping unit. The limit of detection for our method is 5 pmol NO in aqueous solution, which translates into 0.25 nmol L−1 when using a 20 mL seawater sample volume. Our method was applied to measure high-resolution depth profiles of dissolved NO during a cruise to the eastern tropical South Pacific Ocean. It is fast and comparably easy to handle; thus it opens the door for investigating the distribution of NO in the ocean, and it facilitates laboratory studies on NO pathways.

Circulation, eddies, oxygen, and nutrient changes in the eastern tropical South Pacific Ocean

Abstract. A large subsurface oxygen deficiency zone is located in the eastern tropical South Pacific Ocean (ETSP). The large-scale circulation in the eastern equatorial Pacific and off the coast of Peru in November/December 2012 shows the influence of the equatorial current system, the eastern boundary currents, and the northern reaches of the subtropical gyre. In November 2012 the equatorial undercurrent (EUC) is centered at 250 m depth, deeper than in earlier observations. In December 2012, the equatorial water is transported southeastward near the shelf in the Peru–Chile undercurrent (PCUC) with a mean transport of 1.4 Sv. In the oxygen minimum zone (OMZ), the flow is overlaid with strong eddy activity on the poleward side of the OMZ. Floats with parking depth at 400 m show fast westward flow in the mid-depth equatorial channel and sluggish flow in the OMZ. Floats with oxygen sensors clearly show the passage of eddies with oxygen anomalies. The long-term float observations in the upper ocean lead to a net community production estimate at about 18° S of up to 16.7 mmol C m−3 yr−1 extrapolated to an annual rate and 7.7 mmol C m−3 yr−1 for the time period below the mixed layer. Oxygen differences between repeated ship sections are influenced by the Interdecadal Pacific Oscillation (IPO), by the phase of El Niño, by seasonal changes, and by eddies, and hence have to be interpreted with care. At and south of the Equator the decrease in oxygen in the upper ocean since 1976 is related to an increase in nitrate, phosphate, and in part silicate.

Aphotic N2 Fixation in the Eastern Tropical South Pacific Ocean

We examined rates of N2 fixation from the surface to 2000 m depth in the Eastern Tropical South Pacific (ETSP) during El Niño (2010) and La Niña (2011). Replicated vertical profiles performed under oxygen-free conditions show that N2 fixation takes place both in euphotic and aphotic waters, with rates reaching 155 to 509 µmol N m−2 d−1 in 2010 and 24±14 to 118±87 µmol N m−2 d−1 in 2011. In the aphotic layers, volumetric N2 fixation rates were relatively low (<1.00 nmol N L−1 d−1), but when integrated over the whole aphotic layer, they accounted for 87–90% of total rates (euphotic+aphotic) for the two cruises. Phylogenetic studies performed in microcosms experiments confirm the presence of diazotrophs in the deep waters of the Oxygen Minimum Zone (OMZ), which were comprised of non-cyanobacterial diazotrophs affiliated with nifH clusters 1K (predominantly comprised of α-proteobacteria), 1G (predominantly comprised of γ-proteobacteria), and 3 (sulfate reducing genera of the δ-proteobacteria and Clostridium spp., Vibrio spp.). Organic and inorganic nutrient addition bioassays revealed that amino acids significantly stimulated N2 fixation in the core of the OMZ at all stations tested and as did simple carbohydrates at stations located nearest the coast of Peru/Chile. The episodic supply of these substrates from upper layers are hypothesized to explain the observed variability of N2 fixation in the ETSP.

Bacterial transformations of inorganic nitrogen in the oxygen-deficient waters of the Eastern Tropical South Pacific Ocean

Rates of transformations of inorganic nitrogen were measured in the low oxygen, subsurface waters (50–450 m) of the Eastern Tropical South Pacific during February 1985, using 15N tracer techniques. Oxygen concentrations over the entire region were in a range (O 2 < 2.5 μM) that allowed both oxidation and reduction of nitrogen to occur. A wide range of rates was observed for the lowest oxygen levels, indicating that observed oxygen concentration was not a primary factor regulating nitrogen metabolism. High values for subsurface metabolic rates correspond with high levels for surface primary production, both apparently associated with mesoscale features observed in satellite imagery and with mesoscale features of the current field. Measured rates of nitrate reduction and estimated rates of denitrification were sufficient to respire nearly all of the surface primary production that might be transported into the oxygen deficient zone. These results imply that the supply of labile organic material, especially from the surface, was more important than oxygen concentration in modulating the rates of nitrogen transformations within the low oxygen water mass of the Eastern Tropical South Pacific. The pattern of nitrite oxidation and nitrite reduction activities in the oxygen minimum zone supports the hypothesis ( Anderson et al., 1982, Deep-Sea Research, 29, 1113–1140) that nitrite, produced from nitrate reduction, can be recycled by oxidation at the interface between low and high oxygen waters. Rates for denitrification, estimated from nitrate reduction rates, were in harmony with previous estimates based on electron transport system (ETS) measurements and analysis of the nitrate deficit and water residence times. Assimilation rates of NH 4 + were substantial, providing evidence for heterotrophic bacterial growth in low oxygen waters. Ambient concentrations of ammonium were maintained at low values primarily by assimilation; ammonium oxidation was an important mechanism at the surface boundary of the low oxygen zone.

Nitrification, denitrification and nitrous oxide cycling in the eastern tropical South Pacific ocean

A nitrification—denitrification couple that may be summarized by the pathway, NH 4 + → N 2O → N 2 has not been included in past studies of marine denitrification. Neglect of this pathway could lead to underestimates of the amounts of free nitrogen produced by denitrification. Because of this and because our analysis suggests that loss of nitrite from oxygen-deficient waters found in the eastern tropical South Pacific does not introduce major errors, we suggest a total denitrification rate of ∼ 60 Tg N y −1 for the marine water column. Recent studies of the sedimentary denitrification rate also suggest a value of ∼ 60 Tg N y −1, while estimates for the combined nitrogen inputs into the sea via the atmosphere and marine nitrogen fixation have decreased. These findings suggest an unbalanced marine combined-nitrogen budget in which the removal rate significantly exceeds the input. Consideration of the characteristics of nitrifying and denitrifying bacteria and of the distribution of variables in the eastern tropical South Pacific suggest high N 2O turnover rates in eastern boundary regions. Since N 2O may be produced at extremely high rates at the boundary of oxygen-deficient ( O 2 < ∼ 0.1 ml l −1) waters and consumed within these zones, whether the ocean is a source or sink for atmospheric N 2O may depend critically on the exact nature of the dissolved oxygen distribution.