Oxygen Deficient Zone Research Articles

Prolific nitrite reoxidation across the Eastern Tropical North Pacific Ocean

AbstractNitrite is a key intermediate during fixed nitrogen loss in the ocean, and it accumulates within marine Oxygen Deficient Zones (ODZ). ODZs are vast subsurface regions where nitrate is the dominant electron acceptor, and these regions host approximately 50% of the fixed nitrogen loss in the world's oceans. Nitrite accumulates in these waters, and recent research has discovered substantial reoxidation of nitrite back to nitrate, a significant process in the global nitrogen cycle. Partitioning between reduction and oxidation determines if marine fixed nitrogen is lost or recycled. Investigations into nitrite oxidation typically rely on results from incubations, which limits the spatiotemporal sampling coverage. Using basin‐scale data, we analyzed the ratios of nutrient regeneration within the three water masses that feed the Eastern Tropical North Pacific (ETNP) ODZ. Deviations in the ratios of nutrient regeneration from Redfield stoichiometry indicated prolific nitrite reoxidation across this region. We estimate that 79 ± 7% of the nitrite produced in the ODZ between the 26.2 and 26.4 kg m−3 isopycnals is reoxidized, whereas 54 ± 2% of the nitrite produced between the 26.7 and 26.9 kg m−3 isopycnals is reoxidized. Our analysis also illustrates discrete “metabolic switching points” from primarily aerobic to primary anaerobic processes, which occur in each water mass. We applied water mass analysis to repeat cruises on the WOCE P18 line from Baja California to 10°N, which revealed high spatiotemporal variability in nitrite reoxidation. These results confirm previous measurements of significant fixed nitrogen recycling across the ETNP; however, our analysis enables high‐resolution estimates of this process.

Autonomous observations of biogenic N2 in the Eastern Tropical North Pacific using profiling floats equipped with gas tension devices

Oxygen Deficient Zones (ODZs) of the world’s oceans represent a relatively small fraction of the ocean by volume (<0.05% for suboxic and<5% for hypoxic) yet are receiving increased attention by experimentalists and modelers due to their importance in ocean nutrient cycling and predicted susceptibility to expansion and/or contraction forced by global warming. Conventional methods to study these biogeochemically important regions of the ocean have relied on well-developed but still relatively high cost and labor-intensive shipboard methods that include mass-spectrometric analysis of nitrogen-to-argon ratios (N2/Ar) and nutrient stoichiometry (relative abundance of nitrate, nitrite, and phosphate). Experimental studies of denitrification rates and processes typically involve either in-situ or in-vitro incubations using isotopically labeled nutrients. Over the last several years we have been developing a Gas Tension Device (GTD) to study ODZ denitrification including deployment in the largest ODZ, the Eastern Tropical North Pacific (ETNP). The GTD measures total dissolved gas pressure from which dissolved N2 concentration is calculated. Data from two cruises passing through the core of the ETNP near 17 °N in late 2020 and 2021 are presented, with additional comparisons at 12 °N for GTDs mounted on a rosette/CTD as well as modified profiling Argo-style floats. Gas tension was measured on the float with an accuracy of< 0.1% and relatively low precision (< 0.12%) when shallow (P< 200 dbar) and high precision (< 0.03%) when deep (P > 300 dbar). We discriminate biologically produced N2 (ie., denitrification) from N2 in excess of saturation due to physical processes (e.g., mixing) using a new tracer – ‘preformed excess-N2’. We used inert dissolved argon (Ar) to help test the assumption that preformed excess-N2 is indeed conservative. We used the shipboard measurements to quantify preformed excess-N2 by cross-calibrating the gas tension method to the nutrient-deficit method. At 17 °N preformed excess-N2 decreased from approximately 28 to 12 µmol/kg over σ0 = 24–27 kg/m3 with a resulting precision of ±1 µmol N2/kg; at 12 °N values were similar except in the potential density range of 25.7< σ0< 26.3 where they were lower by 1 µmol N2/kg due likely to being composed of different source waters. We then applied these results to gas tension and O2 (< 3 µmol O2/kg) profiles measured by the nearby float to obtain the first autonomous biogenic N2 profile in the open ocean with an RMSE of ± 0.78 µM N2, or ± 19%. We also assessed the potential of the method to measure denitrification rates directly from the accumulation of biogenic N2 during the float drifts between profiling. The results suggest biogenic N2 rates of ±20 nM N2/day could be detected over >16 days (positive rates would indicate denitrification processes whereas negative rates would indicate predominantly dilution by mixing). These new observations demonstrate the potential of the gas tension method to determine biogenic N2 accurately and precisely in future studies of ODZs.

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.

Associations between picocyanobacterial ecotypes and cyanophage host genes across ocean basins and depth.

Cyanophages, viruses that infect cyanobacteria, are globally abundant in the ocean's euphotic zone and are a potentially important cause of mortality for marine picocyanobacteria. Viral host genes are thought to increase viral fitness by either increasing numbers of genes for synthesizing nucleotides for virus replication, or by mitigating direct stresses imposed by the environment. The encoding of host genes in viral genomes through horizontal gene transfer is a form of evolution that links viruses, hosts, and the environment. We previously examined depth profiles of the proportion of cyanophage containing various host genes in the Eastern Tropical North Pacific Oxygen Deficient Zone (ODZ) and at the subtropical North Atlantic (BATS). However, cyanophage host genes have not been previously examined in environmental depth profiles across the oceans. We examined geographical and depth distributions of picocyanobacterial ecotypes, cyanophage, and their viral-host genes across ocean basins including the North Atlantic, Mediterranean Sea, North Pacific, South Pacific, and Eastern Tropical North and South Pacific ODZs using phylogenetic metagenomic read placement. We determined the proportion of myo and podo-cyanophage containing a range of host genes by comparing to cyanophage single copy core gene terminase (terL). With this large dataset (22 stations), network analysis identified statistical links between 12 of the 14 cyanophage host genes examined here with their picocyanobacteria host ecotypes. Picyanobacterial ecotypes, and the composition and proportion of cyanophage host genes, shifted dramatically and predictably with depth. For most of the cyanophage host genes examined here, we found that the composition of host ecotypes predicted the proportion of viral host genes harbored by the cyanophage community. Terminase is too conserved to illuminate the myo-cyanophage community structure. Cyanophage cobS was present in almost all myo-cyanophage and did not vary in proportion with depth. We used the composition of cobS phylotypes to track changes in myo-cyanophage composition. Picocyanobacteria ecotypes shift with changes in light, temperature, and oxygen and many common cyanophage host genes shift concomitantly. However, cyanophage phosphate transporter gene pstS appeared to instead vary with ocean basin and was most abundant in low phosphate regions. Abundances of cyanophage host genes related to nutrient acquisition may diverge from host ecotype constraints as the same host can live in varying nutrient concentrations. Myo-cyanophage community in the anoxic ODZ had reduced diversity. By comparison to the oxic ocean, we can see which cyanophage host genes are especially abundant (nirA, nirC, and purS) or not abundant (myo psbA) in ODZs, highlighting both the stability of conditions in the ODZ and the importance of nitrite as an N source to ODZ endemic LLV Prochlorococcus.

Coincident Biogenic Nitrite and pH Maxima Arise in the Upper Anoxic Layer in the Eastern Tropical North Pacific

AbstractThe Eastern Tropical North Pacific (ETNP), like the other marine oxygen deficient zones (ODZs), is characterized by an anoxic water column, nitrite accumulation at the anoxic core, and fixed nitrogen loss via nitrite reduction to N2O and N2 gases. Here, we constrain the relative contribution of biogeochemical processes to observable features such as the secondary nitrite maximum (SNM) and local pH maximum by simultaneous measurement of inorganic nitrogen and carbon species. High‐resolution sampling within the top 1 km of the water column reveals consistent chemical features previously unobserved in the region, including a tertiary nitrite maximum. Dissolved inorganic carbon measurements show that pH increases with depth at the top of the ODZ, peaking at the potential density of the SNM at σθ = 26.15 ± 0.06 (1 s.d.). We developed a novel method to determine the relative contributions of anaerobic ammonium oxidation (anammox), denitrification, nitrite oxidation, dissimilatory nitrate reduction to nitrite, and calcium carbonate dissolution to the nitrite cycling in the anoxic ODZ core. The calculated relative contributions of each reaction are slightly sensitive to the assumed C:N:P ratio and the carbon oxidation state of the organic matter sinking through the ODZ. Furthermore, we identify the source of the pH increase at the top of ODZ as the net consumption of protons via nitrite reduction to N2 by the denitrification process. The increase in pH due to denitrification impacts the buffering effect of calcite and aragonite dissolving in the ETNP.

Oceanic nutrient rise and the late Miocene inception of Pacific oxygen-deficient zones

The modern Pacific Ocean hosts the largest oxygen-deficient zones (ODZs), where oxygen concentrations are so low that nitrate is used to respire organic matter. The history of the ODZs may offer key insights into ocean deoxygenation under future global warming. In a 12-My record from the southeastern Pacific, we observe a >10‰ increase in foraminifera-bound nitrogen isotopes (15N/14N) since the late Miocene (8 to 9 Mya), indicating large ODZs expansion. Coinciding with this change, we find a major increase in the nutrient content of the ocean, reconstructed from phosphorus and iron measurements of hydrothermal sediments at the same site. Whereas global warming studies cast seawater oxygen concentrations as mainly dependent on climate and ocean circulation, our findings indicate that modern ODZs are underpinned by historically high concentrations of seawater phosphate.

Expression Levels of hgcAB Genes and Mercury Availability Jointly Explain Methylmercury Formation in Stratified Brackish Waters.

Neurotoxic methylmercury (MeHg) is formed by microbial methylation of inorganic divalent Hg (HgII) and constitutes severe environmental and human health risks. The methylation is enabled by hgcA and hgcB genes, but it is not known if the associated molecular-level processes are rate-limiting or enable accurate prediction of MeHg formation in nature. In this study, we investigated the relationships between hgc genes and MeHg across redox-stratified water columns in the brackish Baltic Sea. We showed, for the first time, that hgc transcript abundance and the concentration of dissolved HgII-sulfide species were strong predictors of both the HgII methylation rate and MeHg concentration, implying their roles as principal joint drivers of MeHg formation in these systems. Additionally, we characterized the metabolic capacities of hgc+ microorganisms by reconstructing their genomes from metagenomes (i.e., hgc+ MAGs), which highlighted the versatility of putative HgII methylators in the water column of the Baltic Sea. In establishing relationships between hgc transcripts and the HgII methylation rate, we advance the fundamental understanding of mechanistic principles governing MeHg formation in nature and enable refined predictions of MeHg levels in coastal seas in response to the accelerating spread of oxygen-deficient zones.

Enhanced ocean oxygenation during Cenozoic warm periods

Dissolved oxygen (O2) is essential for most ocean ecosystems, fuelling organisms’ respiration and facilitating the cycling of carbon and nutrients. Oxygen measurements have been interpreted to indicate that the ocean’s oxygen-deficient zones (ODZs) are expanding under global warming1,2. However, models provide an unclear picture of future ODZ change in both the near term and the long term3–6. The paleoclimate record can help explore the possible range of ODZ changes in warmer-than-modern periods. Here we use foraminifera-bound nitrogen (N) isotopes to show that water-column denitrification in the eastern tropical North Pacific was greatly reduced during the Middle Miocene Climatic Optimum (MMCO) and the Early Eocene Climatic Optimum (EECO). Because denitrification is restricted to oxygen-poor waters, our results indicate that, in these two Cenozoic periods of sustained warmth, ODZs were contracted, not expanded. ODZ contraction may have arisen from a decrease in upwelling-fuelled biological productivity in the tropical Pacific, which would have reduced oxygen demand in the subsurface. Alternatively, invigoration of deep-water ventilation by the Southern Ocean may have weakened the ocean’s ‘biological carbon pump’, which would have increased deep-ocean oxygen. The mechanism at play would have determined whether the ODZ contractions occurred in step with the warming or took centuries or millennia to develop. Thus, although our results from the Cenozoic do not necessarily apply to the near-term future, they might imply that global warming may eventually cause ODZ contraction.

Millennial-scale marine productivity changes and the persistence of Antarctic intermediate water off Baja California, Mexico, during the last 6–60 kyr

A high-resolution record of δ18O and δ13C of Uvigerina peregrina (hereinafter: δ18OU and δ13CU) and biogenic opal was used to reconstruct water column conditions in the SW margin of the Baja California peninsula for the last 6–60 kyr. The core was collected from a depth of 700 m in the oxygen deficient zone of the Magdalena margin. Although the δ18OU values showed a large change associated with the passage from glacial conditions to the Holocene, the MIS-3 and MIS-2 were characterized by a low variability. Our results indicated a rapid cooling and/or increase in salinity immediately after H5 that persisted until H1. However, the δ13CU values showed significant variability associated to the incursions of the Antarctic Intermediate Water (AAIW) in the Magdalena margin during the MIS-3 and MIS-2. This agrees with the δ13C composition of water mass end-members at intermediate depths in the Pacific Ocean. Enhanced ventilation during MIS-3 was inferred from the δ13CU values, which were very similar to the δ13C values of the AAIW end-members evidencing the persistence of the AAIW in the region. This conclusion is also supported by the trend of sedimentary δ15N and other dissolved oxygen proxies in the area. A depletion of the δ13CU at a millennial scale suggests intermittent pulses of a water mass richer in nutrients, which generated an increase in exported productivity, and thus, concomitantly depleted the dissolved oxygen levels at intermediate depths. Moreover, exported productivity showed a decreasing long-term trend during the second half of MIS-3, briefly punctuated by millennial-scale increases in exported productivity sustained by nutrient export from the Southern Ocean via the AAIW.

Iron and manganese accumulation within the Eastern Tropical North Pacific oxygen deficient zone

The Eastern Tropical North Pacific (ETNP) contains the largest oxygen deficient zone (ODZ) in the modern ocean. We determined dissolved concentrations of Fe, Fe(II), and Mn from three cruises in the region. Similar to other reported ODZs, Fe(II) was highest in the depth range associated with the secondary nitrite maximum. The main source of this feature is likely lateral advection of water overlying reducing shelf sediments within a narrow density range centered on the potential density anomaly of 26.5 kg/m3. This density horizon is similar to the Eastern Tropical South Pacific (ETSP) and reflects the intersection of the same density range with a large fraction of the continental shelf bottom waters. We also observed subsurface maxima of dissolved Mn in this density range, in contrast to the ETSP. Deep waters were enriched in Fe within the ETNP, analogous to other eastern boundary upwelling systems as well as the Arabian Sea. We argue that in these systems, reducing conditions on the shelf and overlying water column facilitate a robust shelf to basin shuttle of Fe, moving Fe from the coastal margin to deep plumes. Mn is also transported offshore in the core of the ODZ, and the relationship between Fe(II), Mn, and nitrite is remarkably similar between the ETNP, ETSP, and Arabian Sea. The exception is that Mn supply from the Peruvian shelf is less pronounced than in the other two ODZs, potentially reflecting the absence of large rivers in the Peruvian system.

Identifying the Sources and Drivers of Nitrous Oxide Accumulation in the Eddy‐Influenced Eastern Tropical North Pacific Oxygen‐Deficient Zone

AbstractNitrous oxide (N2O) is a powerful greenhouse gas, and oceanic sources account for up to one third of the total natural flux to the atmosphere. In oxygen‐deficient zones (ODZs) like the Eastern Tropical North Pacific (ETNP), N2O can be produced and consumed by several biological processes. In this study, the concentration and isotopocule ratios of N2O from a 2016 cruise in the ETNP were analyzed to examine sources of and controls on N2O cycling across this region. Along the north‐south transect, three distinct biogeochemical regimes were identified: background, core‐ODZ, and high‐N2O stations. Background stations were characterized by smaller variations in N2O concentration and isotopic profiles relative to the other regimes. Core‐ODZ stations were characterized by co‐occurring N2O production and consumption at anoxic depths, indicated by high δ18O‐N2O (>90‰) and low δ15N2Oβ (<−10‰) values, and confirmed by a time‐dependent model, which indicated that N2O production via denitrification was significant and may occur with a nonzero site preference. High‐N2O stations, located at the periphery of a mesoscale eddy, were defined by N2O reaching 126.07 ± 12.6 nM and low oxygen concentrations expanding into near‐surface isopycnals. At these stations, model results indicated significant N2O production from ammonia‐oxidizing archaea and denitrification from nitrate at the N2O maximum within the oxycline, while bacterial nitrification and denitrification from nitrite were insignificant. This study also represents the first in the ETNP to link N2O production mechanisms to a mesoscale eddy through isotopocule measurements, suggesting the importance of eddies to spatiotemporal variability in N2O cycling and emissions from this region.

Two Metatranscriptomic Profiles through Low-Dissolved-Oxygen Waters (DO, 0 to 33 µM) in the Eastern Tropical North Pacific Ocean.

ABSTRACTWe present 16 seawater metatranscriptomes collected from a marine oxygen-deficient zone (ODZ) in the eastern tropical North Pacific (ETNP). This data set will be useful for identifying shifts in microbial community structure and function through oxic/anoxic transition zones, where overlapping aerobic and anaerobic microbial processes impact marine biogeochemical cycling.

An analysis of protists in Pacific oxygen deficient zones: implications for Prochlorococcus and N2 -producing bacteria.

Ocean oxygen deficient zones (ODZs) host 30%-50% of marine N2 production. Cyanobacteria photosynthesizing in the ODZ create a secondary chlorophyll maximum and provide organic matter to N2 -producing bacteria. This chlorophyll maximum is thought to occur due to reduced grazing in anoxic waters. We first examine ODZ protists with long amplicon reads. We then use non-primer-based methods to examine the composition and relative abundance of protists in metagenomes from the Eastern Tropical North and South Pacific ODZs and compare these data to the oxic Hawaii Ocean Time-series (HOT) in the North Pacific. We identify and quantify protists in proportion to the total microbial community. From metagenomic data, we see a large drop in abundance of fungi and protists such as choanoflagellates, radiolarians, cercozoa and ciliates in the ODZs but not in the oxic mesopelagic at HOT. Diplonemid euglenozoa were the only protists that increased in the ODZ. Dinoflagellates and foraminifera reads were also present in the ODZ though less abundant compared to oxic waters. Denitrification has been found in foraminifera but not yet in dinoflagellates. DNA techniques cannot separate dinoflagellate cells and cysts. Metagenomic analysis found taxonomic groups missed by amplicon sequencing and identified trends in abundance.

Bacteriohopanetetrol-<i>x</i>: constraining its application as a lipid biomarker for marine anammox using the water column oxygen gradient of the Benguela upwelling system

Abstract. Interpreting lipid biomarkers in the sediment archive requires a good understanding of their application and limitations in modern systems. Recently it was discovered that marine bacteria performing anaerobic ammonium oxidation (anammox), belonging to the genus Ca. Scalindua, uniquely synthesize a stereoisomer of bacteriohopanetetrol (“BHT-x”). The ratio of BHT-x over total bacteriohopanetetrol (BHT, ubiquitously synthesized by diverse bacteria) has been suggested as a proxy for water column anoxia. As BHT has been found in sediments over 50 Myr old, BHT-x has the potential to complement and extend the sedimentary biomarker record of marine anammox, conventionally constructed using ladderane lipids. Yet, little is known about the distribution of BHT-x in relation to the distribution of ladderanes and to the genetic evidence of Ca. Scalindua in modern marine systems. Here, we investigate the distribution of BHT-x and the application of the BHT-x ratio in relation to distributions of ladderane intact polar lipids (IPLs), ladderane fatty acids (FAs) and Ca. Scalindua 16S rRNA genes in suspended particulate matter (SPM) from the water column of the Benguela upwelling system (BUS), sampled across a large oxygen gradient. In BUS SPM, high BHT-x abundances were restricted to the oxygen-deficient zone on the continental shelf (at [O2] < 45 µmol L−1, in all but one case). High BHT-x abundances co-occurred with high abundances of the Ca. Scalindua 16S rRNA gene (relative to the total number of bacterial 16S rRNA genes) and ladderane IPLs. At shelf stations with [O2] > 50 µmol L−1, the BHT-x ratio was < 0.04 (in all but one case). In apparent contradiction, ladderane FAs and low abundances of BHT and BHT-x (resulting in BHT-x ratios > 0.04) were also detected in oxygenated offshore waters ([O2] up to 180 µmol L−1), whereas ladderane IPLs were undetected. The index of ladderane lipids with five cyclobutane rings (NL5) correlates with in situ temperature. NL5-derived temperatures suggested that ladderane FAs in the offshore waters were not synthesized in situ but were transported down-slope from warmer shelf waters. Thus, in sedimentary archives of systems with known lateral organic matter transport, such as the BUS, relative BHT and BHT-x abundances should be carefully considered. In such systems, a higher BHT-x ratio may act as a safer threshold for deoxygenation and/or Ca. Scalindua presence: our results and previous studies indicate that a BHT-x ratio of ≥ 0.2 is a robust threshold for oxygen-depleted waters ([O2] < 50 µmol kg−1). In our data, ratios of ≥ 0.2 coincided with Ca. Scalindua 16S rRNA genes in all samples (n=62), except one. Lastly, when investigating in situ anammox, we highlight the importance of using ladderane IPLs over BHT-x and/or ladderane FAs; these latter compounds are more recalcitrant and may derive from transported fossil anammox bacteria remnants.

Protein cycling in the eastern tropical North Pacific oxygen‐deficient zone: A de novo‐discovery peptidomic approach

AbstractPeptides are identified using a de novo‐discovery approach in suspended and sinking particles from the eastern tropical North Pacific oxygen‐deficient zone (ODZ) and in a culture of a dominant autotroph from the region, the cyanobacterium Prochlorococcus. The benchmarking experiment with Prochlorococcus shows de novo peptides to be taxonomically specific, and thus of value in augmenting database‐driven approaches. Analysis of the suspended and sinking particles using the de novo‐discovery approach reveals the presence of fungal proteins in deep sinking particles that were not in our original search database, contributing to growing recognition that fungi may play important roles in marine organic matter cycling. Cyanobacterial peptides that have been post‐translationally modified were tracked to depth, where they contribute ~ 1% of the phylum‐level identifiable peptide pool in the sediment trap sample. The majority of peptides found at depth in the detrital pool are associated with membranes, indicating that cellular location is associated with early preservation within the detrital pool. Modified amino acids in sinking and suspended particles include high levels of deamidation, suggesting that partial extracellular degradation of protein could fuel observed anammox and contribute to pools of refractory organic nitrogen.

Slow Particle Remineralization, Rather Than Suppressed Disaggregation, Drives Efficient Flux Transfer Through the Eastern Tropical North Pacific Oxygen Deficient Zone

AbstractModels and observations suggest that particle flux attenuation is lower across the mesopelagic zone of anoxic environments compared to oxic environments. Flux attenuation is controlled by microbial metabolism as well as aggregation and disaggregation by zooplankton, all of which shape the relative abundance of differently sized particles. Observing and modeling particle spectra can provide information about the contributions of these processes. We measured particle size spectrum profiles at one station in the oligotrophic Eastern Tropical North Pacific Oxygen Deficient Zone (ETNP ODZ) using an underwater vision profiler (UVP), a high‐resolution camera that counts and sizes particles. Measurements were taken at different times of day, over the course of a week. Comparing these data to particle flux measurements from sediment traps collected over the same time‐period allowed us to constrain the particle size to flux relationship, and to generate highly resolved depth and time estimates of particle flux rates. We found that particle flux attenuated very little throughout the anoxic water column, and at some time points appeared to increase. Comparing our observations to model predictions suggested that particles of all sizes remineralize more slowly in the ODZ than in oxic waters, and that large particles disaggregate into smaller particles, primarily between the base of the photic zone and 500 m. Acoustic measurements of multiple size classes of organisms suggested that many organisms migrated, during the day, to the region with high particle disaggregation. Our data suggest that diel‐migrating organisms both actively transport biomass and disaggregate particles in the ODZ core.

Microbial Eukaryotes Associated With Sediments in Deep-Sea Cold Seeps.

Microbial eukaryotes are key components of the marine food web, but their distribution in deep-sea chemosynthetic ecosystems has not been well studied. Here, high-throughput sequencing of the 18S rRNA gene and network analysis were applied to investigate the diversity, distribution and potential relationships between microbial eukaryotes in samples collected from two cold seeps and one trough in the northern South China Sea. SAR (i.e., Stramenopiles, Alveolata, and Rhizaria) was the predominant group in all the samples, and it was highly affiliated to genotypes with potential symbiotic and parasitic strategies identified from other deep-sea extreme environments (e.g., oxygen deficient zones, bathypelagic waters, and hydrothermal vents). Our findings indicated that specialized lineages of deep-sea microbial eukaryotes exist in chemosynthetic cold seeps, where microbial eukaryotes affiliated with parasitic/symbiotic taxa were prevalent in the community. The biogeographic pattern of the total community was best represented by the intermediate operational taxonomic unit (OTU) category, whose relative abundance ranged 0.01–1% within a sample, and the communities of the two cold seeps were distinct from the trough, which suggests that geographical proximity has no critical impact on the distribution of deep-sea microbial eukaryotes. Overall, this study has laid the foundations for future investigations regarding the ecological function and in situ trophic relationships of microbial eukaryotes in deep-sea ecosystems.

A High‐Resolution Atlas of the Eastern Tropical Pacific Oxygen Deficient Zones

AbstractOxygen deficient zones (ODZs) are important biogeochemical provinces of the global oceans wherein standing dissolved oxygen concentrations decrease to nanomolar levels. Despite their confinement, these regions are disproportionally important to the ocean's role in modulating Earth's climate through the interactions between the marine nitrogen cycle and that of carbon. Moreover, the spatial domain of low oxygen regions of the ocean is predicted to change as a consequence of ocean warming, increased stratification, and changes in circulation and productivity. However, the expanse of the modern ODZs is poorly resolved due to a dearth of direct sampling compounded with errors that arise in the processing and gridding of the sparse measurements that do exist. Here, we take a novel approach to map the horizontal and vertical extent of the two major ODZs of the eastern tropical Pacific via analysis of meter‐scale resolution electrode sensors from both ship casts and Argo profiles, rather than from discretized bottle measurements. The resulting three‐dimensional data product is based on a compendium of nearly 15 million measurements taken across three decades and provides the precise locations of low oxygen water, elucidating the ODZs' three‐dimensional structures. It can be utilized by researchers to validate models, plan cruise occupations, and as a comparison for future change. Calculations made with this high‐resolution atlas also provide the volumes, layers of maximal areal extent, and other descriptive statistics for both Pacific ODZs. Finally, the atlas reveals fine‐scale features of oxygenated water mass intrusions and regional differences across these anoxic zones.

Untangling the effect of organic matter and dissolved oxygen on living benthic foraminifera in the southeastern Arabian Sea

We studied the relationship between organic carbon in sediments (%Corg), bottom water dissolved oxygen and living benthic foraminifera in a marginal marine upwelling setting from the southeastern Arabian Sea to develop proxy indicator for marine productivity as well as oxygen deficient zones. The surface sediments from 43 stations covering a depth range of 25 to 2980 m were used. The relationship between living benthic foraminifera and ambient environmental parameters (seawater temperature, salinity, dissolved oxygen, organic carbon and Corg/TN and depth) was assessed by multivariate analysis. A majority of the living benthic foraminifera were significantly affected by several ambient parameters. We report that the living benthic foraminiferal assemblage comprising of Epistominella umbonifera, Uvigerina auberiana, Reophax longicollis and Osangularia bengalensis is significantly affected only by %Corg in the sediment. Additionally, we also found that the assemblage including Bolivina obscuranta, Bulimina arabiensis, Bulimina pseudoaffinis and Cancris penangensis is significantly affected only by the bottom water dissolved oxygen. The living benthic foraminifera assemblages can be used to reconstruct marine productivity and dissolved oxygen concentration in the past.

Organic complexation of iron by strong ligands and siderophores in the eastern tropical North Pacific oxygen deficient zone

Continental margins are an important external source of dissolved iron to the marine environment. However, the mechanisms responsible for the offshore transport of dissolved iron is impacted by the resulting iron speciation. We characterized the iron speciation in the Eastern Tropical North Pacific (ETNP) oxygen deficient zone (ODZ), including dissolved iron, organic iron-binding ligands, and reduced iron. Organic iron-binding ligands were measured using both competitive ligand exchange adsorptive cathodic stripping voltammetry (CLE-ACSV) and liquid chromatography electrospray ionization mass spectrometry (LC-ESI-MS) in order to explore the impact of organic ligands on dissolved iron (dFeT) and iron(II) biogeochemistry in the region. Organic ligands were present in high concentrations (1.06–5.30 nmol L−1) and exceeded dissolved iron concentrations (0.36–4.52 nmol L−1) at all locations. Iron-binding strengths (logKFeL,Fe′cond) ranged 11.22 to 12.75 and were elevated in the ODZ layer relative to the oxygenated water column. LC-ESI-MS revealed the presence of siderophores, or bacterially-produced organic ligands with high Fe-affinity, in all samples analyzed, suggesting these compounds may be produced by microbes in the ODZ despite high ambient dFeT concentrations. This study is the first to characterize siderophores in an ODZ environment to date, and the three siderophores found (amphibactin B, synechobactin c9, synechobactin c10) could contribute to the observed elevated logKFeL,Fe′cond of ligands in the ODZ. Comparative analysis of organic ligand logKFeL,Fe′cond values in other low oxygen environments suggests that strong ligands, including siderophores, could be present in other low oxygen regions. In a simple model of the shelf-to-offshore iron transport mechanism, strong organic iron-binding ligands had a large impact on the longevity and transport of iron in the ODZ. These results suggest that organic ligand composition can have an impact on iron distributions in the ETNP ODZ and regulate the offshore transport of iron to the open ocean.