Oxygen Minimum Zone Research Articles

What Controls the Large‐Scale Efficiency of Carbon Transfer Through the Ocean's Mesopelagic Zone? Insights From a New, Mechanistic Model (MSPACMAM)

AbstractA key challenge for current‐generation Earth system models (ESMs) is the simulation of the penetration of sinking particulate organic carbon (POC) into the ocean interior, which has implications for projections of future oceanic carbon sequestration in a warming climate. This paper presents a new, cost‐efficient, mechanistic 1D model that prognostically calculates POC fluxes by carrying four component particles in two different size classes. Gravitational settling and removal/transformation processes are represented explicitly through parameterizations that incorporate the effects of particle size and density, dissolved oxygen, calcite and aragonite saturation states, and seawater temperature, density, and viscosity. The model reproduces the observed POC flux attenuation at 22 locations in the North Atlantic and North Pacific. The model is applied over a global ocean domain with seawater properties prescribed from observation‐based climatologies in order to address an important scientific question: What controls the spatial pattern of mesopelagic POC transfer efficiency? The simulated vertical POC transfer is more efficient at high latitudes than at low latitudes with the exception of oxygen minimum zones, which is consistent with recent inverse modeling and neutrally buoyant sediment trap studies. Here, model experiments show that the relative abundance of large‐sized, rapidly sinking particles and the slower rate of remineralization at high latitudes compensate for the region's lack of calcium carbonate ballast and the cold‐water viscous resistance, leading to higher transfer efficiencies compared to low‐latitude regions. The model could be deployed in ESMs in order to diagnose the impacts of climate change on oceanic carbon sequestration and vice versa.

Elevated Mercury Concentrations and Isotope Signatures (N, C, Hg) in Yellowfin Tuna (Thunnus albacares) from the Galápagos Marine Reserve and Waters off Ecuador

We examined how dietary factors recorded by C and N influence Hg uptake in 347 individuals of yellowfin tuna (Thunnus albacares), an important subsistence resource from the Galápagos Marine Reserve (Ecuador) and the Ecuadorian mainland coast in 2015-2016. We found no differences in total Hg (THg) measured in red muscle between the two regions and no seasonal differences, likely due to the age of the fish and slow elimination rates of Hg. Our THg concentrations are comparable to those of other studies in the Pacific (0.20-9.60 mg/kg wet wt), but a subset of individuals exhibited the highest Hg concentrations yet reported in yellowfin tuna. Mercury isotope values differed between Δ199 Hg and δ202Hg in both regions (Δ199 Hg = 2.86 ± 0.04‰ vs. Δ199 Hg = 2.33 ± 0.07‰), likely related to shifting food webs and differing photochemical processing of Hg prior to entry into the food web. There were significantly lower values of both δ15 N and δ13 C in tuna from Galápagos Marine Reserve (δ15 N: 8.5-14.2‰, δ13 C: -18.5 to -16.1‰) compared with those from the Ecuadorian mainland coast (δ15 N: 8.3-14.4‰, δ13 C: -19.4 to -11.9‰), of which δ13 C values suggest spatially constrained movements of tuna. Results from the pooled analysis, without considering region, indicated that variations in δ13 C and δ15 N values tracked changes of Hg stable isotopes. Our data indicate that the individual tuna we used were resident fish of each region and were heavily influenced by upwellings related to the eastern Pacific oxygen minimum zone and the Humboldt Current System. The isotopes C, N, and Hg reflect foraging behavior mainly on epipelagic prey in shallow waters and that food web shifts drive Hg variations between these populations of tuna. Environ Toxicol Chem 2022;41:2732-2744. © 2022 SETAC.

Bacterial community of sediments under the Eastern Boundary Current System shows high microdiversity and a latitudinal spatial pattern

The sediments under the Oxygen Minimum Zone of the Eastern Boundary Current System (EBCS) along Central-South Peru and North-Central Chile, known as Humboldt Sulfuretum (HS), is an organic-matter-rich benthic habitat, where bacteria process a variety of sulfur compounds under low dissolved-oxygen concentrations, and high sulfide and nitrate levels. This study addressed the structure, diversity and spatial distribution patterns of the HS bacterial community along Northern and South-Central Chile using 16S rRNA gene amplicon sequencing. The results show that during the field study period, the community was dominated by sulfur-associated bacteria. Indeed, the most abundant phylum was Desulfobacterota, while Sva0081 sedimentary group, of the family Desulfosarcinaceae (the most abundant family), which includes sulfate-reducer and H2 scavenger bacteria, was the most abundant genus. Furthermore, a spatial pattern was unveiled along the study area to which the family Desulfobulbaceae contributed the most to the spatial variance, which encompasses 42 uncharacterized amplicon sequence variants (ASVs), three assigned to Ca. Electrothrix and two to Desulfobulbus. Moreover, a very high microdiversity was found, since only 3.7% of the ASVs were shared among localities, reflecting a highly diverse and mature community.

Ferromanganese crusts of the Vietnam margin, central South China Sea: Composition and genesis

Ferromanganese (FeMn) crusts collected from two seamounts of the Vietnam continental margin, central South China Sea were analyzed for mineralogical and chemical compositions, and dated using 230Thexcees technique and cobalt chronometry. The studies of dissolved oxygen content in water column at the area by CTD-profiling display Oxygen Minimum Zone locates at the depth 420 to 1100 m with O2 concentration of 1.6–1.8 ml/l. The composition and morphology of studied crusts are different from Prime Crust Zone crusts in Pacific Ocean and partially similar to ferromanganese crusts and nodules from northern South China Sea due to unique environment of basin. The FeMn crusts from Vietnam margin show high ratios of Fe relative to Mn and Ni relative to Co, high Th and Pb concentrations and comparable to PCZ deposits low contents of REE. Few main mechanisms were dominated in crust formation at Vietnam margin including a permanent input of detrital material sourced by river discharge, shelf sediments and dust plume while Fe and Mn (oyxhydr)oxides slow precipitate from seawater as initially colloidal particles within oxygen-minimum zone predominantly at the surface of biogenic limestone substrate. Upwelling and high coastal primary productivity are important factors in Fe, Ni, Th and Pb transportation and accumulation. The main factor of high Fe/Mn ratio is the significant flux of Fe released by redox cycling in the sediment. SCS crusts are enriched in Ni relative to Co due to influx of Ni from coastal laterite weathering and optimum conditions for its transportation to crust formation area. Thorium and lead related to significant continental detrital input and eolian supply, especially Asian dust. Due to diversity of crust thicknesses local hydrographic current regime is also suggested as crucial factor in crust formation. Calculated growth rate using 230Thexcees technique is 11 ± 1 mm/Myr and 4.4 ± 0.5 mm/Myr for two measured crusts. The crust growth in South China Sea might be initiated around 1.5 Myr ago, the period which coincide with the Southern Ocean glacial-induced restructuring dated 1.6 Myr.

Seals, whales and the Cenozoic decline of nautiloid cephalopods

AbstractAimNautilusandAllonautilus, last members of the once widespread nautiloid cephalopods, are today restricted to the deep central Indo‐West Pacific Ocean, for reasons that remain unclear. Cephalopod evolution is generally considered as being driven by vertebrate predation; therefore, we investigated the role of whales and seals in the decline of nautiloids through the Cenozoic.LocationGlobal.TaxonNautiloids, pinnipeds, cetaceans.MethodsDistribution data for nautiloids, pinnipeds and cetaceans through the Cenozoic were compiled and plotted on a series of paleogeographic maps. Nautiloid shell sizes were compiled and plotted against the first appearance of pinnipeds and cetaceans in key regions.ResultsFrom the Oligocene onward, nautiloids became extinct in areas where pinnipeds appeared. The exception is the agile nautiloidAturia, extinct globally at the end of the Miocene. A major role of odontocetes in the demise of nautiloids is not apparent, except for a few brevirostrine Oligocene taxa from the North American Atlantic and Pacific coasts, which appeared in these areas at the same time as nautilids disappeared. The Oligocene disappearance of nautiloids (exceptAturia) from the American Pacific coasts coincides with the development of oxygen minimum zones (OMZs) in this region.Main conclusionsWe hypothesize that the Cenozoic spread of pinnipeds drove nautiloids into their present‐day central Indo‐West Pacific refuge. Additional factors for the local extinction of nautiloids in the Oligocene include predation by short‐snouted whales and the development of OMZs, preventing nautiloids from retreating into deeper water.

Contribution of Carbon Fixation Toward Carbon Sink in the Ocean Twilight Zone

AbstractDark carbon (C) fixation in the ocean twilight zone plays a crucial role toward C sink, but its potential has not been tested sufficiently in experiments. Here we analyzed dark C fixation in the twilight zone of the Arabian Sea along with the primary production measurements in the euphotic zone. The average dark C fixation rates in the suboxic oxygen minimum zone (OMZ) waters were higher than that in the hypoxic OMZ waters, which could be explained by the preferential existence of chemoautotrophic ammonium oxidizers and anammox bacteria owing to NO2− maxima in the suboxic OMZ waters. This study supports a previous hypothesis of significant contribution of dark C fixation to sinking C fluxes in the OMZ of the Arabian Sea. Extrapolation of the measured dark C fixation rates to the global ocean ranged up to 7.4 Pg C y−1; amounting to ∼15% of the global ocean primary production.

Metaomics unveils the contribution of Alteromonas bacteria to carbon cycling in marine oxygen minimum zones

Gammaproteobacteria of the genus Alteromonas are prominent members of pelagic marine microbial communities, playing critical roles in the aerobic degradation of particulate organic matter. Comparative genomic studies of these microorganisms have mainly focused on the metabolic and genomic plasticity of strains isolated primarily from oxygenated environments. In this study, we show that Alteromonas significantly contribute to marine microbial communities from suboxic waters ([O2] < 5 uM) in both the free-living (FL) and particle-attached (PA) fractions, but considerably decrease in abundance in the anoxic waters. The highest proportion of Alteromonas transcripts was found within the secondary fluorescence maximum (SFM) of Oxygen Minimum Zones (OMZs). This metatranscriptomic information suggests an in situ coupling of Alteromonas iron (Fe) and carbon metabolisms, and a relevant role of the glyoxylate cycle across the different layers of the OMZs. This study demonstrates that Alteromonas is an abundant and active member of the OMZ microbial communities, with a potentially significant impact on the carbon cycling in these ecosystems. These results provide valuable environmental evidence to support previous culture-based studies assessing the physiology and ecology of these ubiquitous marine heterotrophs under low-oxygen conditions.

Multi-factor biotopes as a method for detailed site characterization in diverse benthic megafaunal communities and habitats in deep-water off Morro Bay, California

Here we describe the methods and results for biological characterization of the benthos on a previously unexplored area of central California, USA seafloor. We conducted 40 remotely operated vehicle dives from 371 to 1173 m water depth. Seafloor habitats and megafauna (fish and invertebrates) were documented from 46.8 km of seafloor video footage. Our expanded development and analysis of biotopes from quantitative data allowed us to describe detailed biological communities, along with the physical characteristics and habitat associations within the study area. This method provides a framework for potential monitoring, detection of future environmental change (natural and anthropogenic) and comprehensive comparison to other deep water regions. From 185 h of observational video at 25 sites, nearly 120,000 annotations of organisms, habitat characters, biological detritus and anthropogenic debris were recorded and analyzed. We identified a total of 228 taxa, with 173 of them present on linear quantitative transects. Species richness for transects ranged from 0.04 to 0.28 m-2 (8–55 taxa), with densities ranging from 0.07 to 5.20 ind. m−2. Both were highest on hard substrate with greatest surface area. Densities decreased with depth. Within soft substratum zones was a large field of pockmarks, which are seafloor depressions averaging 175 m in diameter and 5 m in depth. Pockmarks have sometimes been associated with seafloor gas seepage, but here we found no biological evidence of chemosynthetic organisms. No significant differences were found in either density nor species richness at pockmark sites vs. non-pockmark sites. Mud draped greenish-black coarse sand occurred only in low oxygen areas, while hummocky, rugose mud supported somewhat different species than flat mud plains. Seventy percent of the transects occurred inside the oxygen minimum zone. We conclude that high rugosity, slope, and the presence of hard substratum were better predictors of species richness and density than oxygen concentration in this specific study. Abundant biological detritus, in the form of dead and dying pelagic pyrosomes and salps, created a large, presumably ephemeral flux of carbon to the seafloor during the study period.

High-resolution post-release behaviour and recovery periods of two highly prized recreational sportfish: the blue marlin and sailfish

Abstract High recreational catch rates of istiophorid billfishes in the eastern tropical Pacific (ETP) have led to substantial eco-tourism derived economic benefits for the countries in the region, prompting many countries to mandate catch-and-release practices for recreational anglers. Previous estimates of billfish post-release behaviours and recovery periods after these physiologically stressful capture events, however, vary widely depending on the type of tag used. Using high-resolution, multi-sensor biologging tags, we provide a fine-scale, detailed view of the behaviour and recovery periods of blue marlin (Makaira nigricans; n = 9) and sailfish (Istiophorus platypterus, Istiophoridae; n = 9) caught in a typical recreational fishery in the ETP. Angling times ranged from 4 to 90 min, and fish were monitored for periods of 6–70 h after release. Blue marlin showed a characteristic long, deep dive immediately after release, with significantly greater duration associated with longer fight times, a behaviour not typical for sailfish. Diving depths were, however, much shallower than those previously reported for both species due to the shallow thermocline and oxycline present in the ETP. Data from 40 derived metrics from acceleration (i.e. tailbeat period, amplitude, pitch, etc.) and physical parameters (i.e. depth, speed, temperature, oxygen saturation, etc.) used to quantify a recovery period suggest blue marlin and sailfish recover 9.0 ± 3.2 and 4.9 ± 2.8 h after release, respectively. Our high-resolution assessment of post-release behaviour suggests that these billfish are capable of rapid physiological recovery after capture in recreational fisheries, and that catch-and-release practices like those used here can be an effective approach to conserve and sustain billfish populations in the ETP. Predicted climate change caused shallowing of the oxygen minimum zone, however, would increase the vertical habitat compression present in this region, potentially prolonging or inhibiting recovery.

The protist community traces seasonality and mesoscale hydrographic features in the oligotrophic Sargasso Sea

Protists represent the majority of the eukaryotic diversity in the oceans. They have different functions in the marine food web, playing essential roles in the biogeochemical cycles. While the available data is rich in horizontal and temporal coverage, little is known on their vertical structuring, particularly below the photic zone. The present study applies V4 18S rDNA metabarcoding to samples collected over three years in conjunction with the BATS time-series to assess marine protist communities in the epipelagic and mesopelagic zones (0-1000 m). The protist community showed a dynamic seasonality in the epipelagic, responding to hydrographic yearly cycles. Mixotrophic lineages dominated throughout the year. However, autotrophs bloomed during the rapid transition between the winter mixing and the stratified summer, and heterotrophs had their peak at the end of summer, when the base of the thermocline reaches its deepest depth. Below the photic zone, the community, dominated by Rhizaria, is depth-stratified and relatively constant throughout the year, although they followed local hydrographic and biological features such as the oxygen minimum zone. The results suggest a dynamic partitioning of the water column, where the niche vertical position for each community changes throughout the year in the epipelagic, likely depending on nutrient availability, the mixed layer depth, and other hydrographic features. At depth, the protist community closely tracked mesoscale events (eddies), where the communities followed the hydrographic uplift, raising the deeper communities for hundreds of meters, and compressing the communities above.

Variability of particulate organic carbon and assessment of satellite retrieval algorithms over the eastern Arabian Sea.

Particulate organic carbon (POC) and its variability were studied to assess the accuracy of ocean colour retrieval algorithms over the eastern Arabian Sea (EAS) as it controls the carbon sequestration, oxygen minimum zone and biogeochemical (C, N and P) cycles. The seasonality in the physical and biological processes strongly influenced the distribution of POC along the EAS. Higher POC and chlorophyll a (chl a) during the spring inter monsoon (SIM) in the north EAS were due to detrainment bloom. The lower POC:chl a ratios during the winter monsoon (WM) (299.8 ± 190.8) than the SIM (482.1 ± 438.3) were due to the influence of freshly derived organic matter with high nutrient levels. The moderate coefficient of regression values of POC with chl a (R2 = 0.49, N = 59) suggests the importance of dead organic materials in controlling the POC distribution in the EAS. Validation between satellite and in situ POC using the four ocean colour retrieval algorithms showed that the algorithm based on the ratio of remote sensing reflectance (Rrs) performed better (R2 = 0.6, N = 17). It also showed a linear trend of POC with absorption coefficients suggesting it as an optical proxy for the POC retrieval.

Pacific Equatorial Undercurrent: Mean state, sources, and future changes across models

The Equatorial Undercurrent (EUC) stretches across the Pacific, transporting cool waters rich in oxygen and nutrients eastward to one of the most productive regions in the ocean. As an intricate part of the global climate system, EUC dynamics are essential to understanding future climate change but are poorly represented in global coupled climate models. This study examines EUC representation and future changes in the latest generations of the Coupled Model Intercomparison Project (CMIP6 and CMIP5) and an eddy-permitting ocean model. We also examine historical and projected changes in EUC source waters, including the Mindanao Current (MC), New Guinea Coastal Undercurrent (NGCU), and interior thermocline convergence. The circulation features in the models are broadly consistent with observations and ocean reanalyses, but improvements from CMIP5 to CMIP6 are limited. In the future projections, the EUC is enhanced in the western Pacific, with less prominent changes in CMIP6, but more so in the eddy-permitting model. The western Pacific EUC enhancement is likely associated with a wind-driven redirection of waters south of the equator, in which the NGCU boundary flow increases while the interior thermocline convergence decreases. This is superimposed on an overall weakening of the North Pacific subtropical overturning cell, including the MC, interior thermocline convergence, and Ekman divergence. As EUC heat and nutrient composition is linked to its sources, these projected changes have implications for the EUC's role in air–sea feedbacks, nutrient replenishment, and oxygen minimum zone ventilation in the eastern Pacific.

Quantifying the Contribution of Ocean Mesoscale Eddies to Low Oxygen Extreme Events

AbstractOcean mesoscale eddies have been identified as drivers of localized extremely low dissolved oxygen concentration ([O2]) conditions in the subsurface. We employ a global physical‐biogeochemical ocean model at eddy‐permitting resolution to conduct a census of open‐ocean eddies near Eastern Boundary Upwelling Systems adjacent to tropical Oxygen Minimum Zones (OMZs). We track cyclonic and anticyclonic eddies with a surface signature over the period 1992–2018 and isolate their subsurface oxygen characteristics. We identify strongly deoxygenating eddies and quantify their contribution to low [O2] extreme events. Our results show that model simulated low [O2] extreme event frequency is 2–7 times higher in eddies versus non‐eddying locations, with regionally more than half of low [O2] extreme events outside of the permanent OMZs being associated with eddies. Our study highlights the need for further work to investigate the drivers, characteristics and potential ecosystem impacts of low [O2] extreme events.

The Fate of Oxygen in the Ocean and Its Sensitivity to Local Changes in Biological Production

AbstractWe investigate the sensitivity of the oxygen content and true oxygen utilization of key low‐oxygen regions Ω to pointwise changes in biological production. To understand how the combined water and biogenic particle transport controls the sensitivity patterns and the fate of oxygen in the ocean, we develop new relationships that link the steady‐state oxygen content and deficit of Ω to the downstream and upstream oxygen utilization rate (OUR), respectively. We find that the amount of oxygen from Ω that will be lost per unit volume at point r is linked to OUR(r) through the mean oxygen age accumulated in Ω. The geographic sensitivity pattern of the Ω‐integrated oxygen deficit is shaped by where the utilization occurs that causes this deficit. The contribution to the oxygen deficit of Ω from utilization at r is controlled by the mean time that water at r spends in Ω before next ventilation at the surface. We illustrate these relationships and the new transport timescales using a simple steady‐state data‐constrained carbon and oxygen model. We focus on Ω being the global ocean, the Pacific Hypoxic Zone (PHZ, [O2] < 62.5 µM), and the North Pacific oxygen minimum zone. The oxygen deficit of the PHZ is most sensitive where mode and intermediate waters form and where increased organic‐matter production directly increases the PHZ's oxygen demand. The fraction of the local oxygen concentration that will be utilized in respiration is as high as 90% in the PHZ and up to 70% in the water column beneath it.

Physical-chemical factors influencing the vertical distribution of phototrophic pico-nanoplankton in the Oxygen Minimum Zone (OMZ) off Northern Chile: The relative influence of low pH/low O2 conditions

The vertical distribution of phytoplankton is of fundamental importance in the structure, dynamic, and biogeochemical pathways in marine ecosystems. Nevertheless, what are the main factors determining this distribution remains as an open question. Here, we evaluated the relative influence of environmental factors that might control the coexistence and vertical distribution of pico-nanoplankton associated with the OMZ off northern Chile. Our results showed that in the upper layer Synechococcus-like cells were numerically important at all sampling stations. Pico-nano eukaryotes and phototrophic nanoflagellates (PNF) also showed high abundances in the upper layer decreasing in abundance down to the upper oxycline, while only Prochlorococcus showed high abundances under oxycline and within the oxygen-depleted layer. Statistical analyses evidenced that temperature, oxygen, and carbonate chemistry parameters (pH and dissolved inorganic carbon, DIC) influenced significantly the vertical distribution of phototrophic pico-nanoplankton. Additionally, we experimentally-evaluated the combined effect of low pH/low O2 conditions on a nanophytoplankton species, the haptophyte Imantonia sp. Under control conditions (pH = 8.1; O2 = 287.5 μM, light = 169.6 μEm−2s−1), Imantonia sp. in vivo fluorescence increased over fifty times, inducing supersaturated O2 conditions (900 μM) and an increasing pH (8.5), whereas upon an experimental treatment mimicking OMZ conditions (pH = 7.5; O2 = 55.6 μM; light = 169.6 μEm−2s−1), in vivo fluorescence declined dramatically, suggesting that Imantonia sp. did not survive. Although preliminary, our study provides evidence about the role of low pH/low O2 conditions on the vertical distribution of nanophytoplankton, which deserve future attention through both fieldwork and more extended experimental experiences.

Into the Dark: Exploring the Deep Ocean with Single-Virus Genomics.

Single-virus genomics (SVGs) has been successfully applied to ocean surface samples allowing the discovery of widespread dominant viruses overlooked for years by metagenomics, such as the uncultured virus vSAG 37-F6 infecting the ubiquitous Pelagibacter spp. In SVGs, one uncultured virus at a time is sorted from the environmental sample, whole-genome amplified, and sequenced. Here, we have applied SVGs to deep-ocean samples (200–4000 m depth) from global Malaspina and MEDIMAX expeditions, demonstrating the feasibility of this method in deep-ocean samples. A total of 1328 virus-like particles were sorted from the North Atlantic Ocean, the deep Mediterranean Sea, and the Pacific Ocean oxygen minimum zone (OMZ). For this proof of concept, sixty single viruses were selected at random for sequencing. Genome annotation identified 27 of these genomes as bona fide viruses, and detected three auxiliary metabolic genes involved in nucleotide biosynthesis and sugar metabolism. Massive protein profile analysis confirmed that these viruses represented novel viral groups not present in databases. Although they were not previously assembled by viromics, global fragment recruitment analysis showed a conserved profile of relative abundance of these viruses in all analyzed samples spanning different oceans. Altogether, these results reveal the feasibility in using SVGs in this vast environment to unveil the genomes of relevant viruses.

Mercury stable isotopes suggest reduced foraging depth in oxygen minimum zones for blue sharks

Oxygen minimum zones (OMZs) are currently expanding across the global ocean due to climate change, leading to a compression of usable habitat for several marine species. Mercury stable isotope compositions provide a spatially and temporally integrated view of marine predator foraging habitat and its variability with environmental conditions. Here, we analyzed mercury isotopes in blue sharks Prionace glauca from normoxic waters in the northeastern Atlantic and from the world's largest and shallowest OMZ, located in the northeastern Pacific (NEP). Blue sharks from the NEP OMZ area showed higher Δ199Hg values compared to sharks from the northeastern Atlantic, indicating a reduction in foraging depth of approximately 200 m. Our study suggests for the first time that blue shark feeding depth is altered by expanding OMZs and illustrates the use of mercury isotopes to assess the impacts of ocean deoxygenation on the vertical foraging habitat of pelagic predators.

Nitrogenases in Oxygen Minimum Zone Waters

Biological dinitrogen (N2) fixation is the pathway making the large pool of atmospheric N2 available to marine life. Besides direct rate measurements, a common approach to explore the potential for N2 fixation in the ocean is a screening-based targeting the key functional marker gene nifH, coding for a subunit of the nitrogenase reductase. As novel sequencing techniques improved, our understanding of the diversity of marine N2 fixers grew exponentially. However, one aspect of N2 fixation in the ocean is often underexplored, which are the two alternative types of the key enzyme of N2 fixation, the nitrogenase. Altogether there are three isoenzymes, the most common Mo-Fe nitrogenase Nif, the Fe-Fe nitrogenase Anf, and the V-Fe nitrogenase Vnf, which differ regarding their genetic organization, as well as their metal co-enzymes. While Mo is only available in the presence of at least traces of oxygen (O2), V and Fe are available if O2 is absent. Therefore, low O2 and anoxic ocean environments could be an ideal place to explore the diversity of the different isotypes of the nitrogenases. Most phylogenetic studies, however, were only based on the functional marker gene nifH, encoding for a subunit of the Nif nitrogenase, and thus limited in representing the diversity of alternative nitrogenases. Here, we screened metagenomes and -transcriptomes from O2 minimum zones off Peru, from the Bay of Bengal, and the anoxic Saanich Inlet to explore the diversity of genes involved in N2 fixation. We identified genes related to all three nitrogenases, and a generally increased diversity as compared to our previous nifH based on studies from OMZ waters. While we could not confirm gene expression of alternative nitrogenases from our transcriptomic, we detected diazotrophs harboring the genetic potential for alternative nitrogenases. We suggest that alternative nitrogenases may not be used under conditions present in those waters, however, depending on trace metal availability they may become active under future ocean deoxygenation.

Deglacial restructuring of the Eastern equatorial Pacific oxygen minimum zone

Oxygenation in the Eastern Equatorial Pacific is responsive to ongoing climate change in the modern ocean, although whether the region saw a deglacial change in extent or position of the Oxygen Minimum Zone remains poorly constrained. Here, stable isotopes from the shells of an Oxygen Minimum Zone-dwelling planktic foraminifer are used to reassess the position of the mid-water Oxygen Minimum Zone relative to both the thermocline and benthos. Oxygen isotopes record a rapid shoaling of the Oxygen Minimum Zone towards the thermocline associated with Heinrich Stadial 1 and persisting through the deglaciation. Meanwhile, carbon isotope similarities between Oxygen Minimum Zone-dwelling Globorotaloides hexagonus and benthic Cibicidoides wuellerstorfi suggest a shared source water through the deglaciation. Results support a direct role for the Eastern Equatorial Pacific in venting carbon to the atmosphere through the deglaciation, a deglacial expansion of the Oxygen Minimum Zone, and a restructuring of mid-water oxygen and carbon dynamics from the glacial to Holocene intervals.

Microbial production of toluene in oxygen minimum zone waters in the Humboldt Current System off Chile

Expansion of oxygen minimum zones in the world's oceans is likely to enhance the production of anaerobic metabolites by marine microorganisms. Here we show that toluene is present throughout the year in shelf waters of the upwelling ecosystem off Concepción (36° S), Chile, and it is a product of microbial anaerobic metabolism. The intra-annual variability in toluene concentrations is consistent with seasonal variability in the strengths of suboxic equatorial and oxygenated subantarctic water masses. Laboratory incubations of oxygen minimum zone water showed microbial production of toluene in the absence of O2. Toluene concentrations were elevated (up to 96 nM) in deeper O2-depleted waters and followed a seasonal pattern in oceanographic conditions. There is evidence to hypothesize that microbial production of toluene could be a homeostatic biochemical mechanism to thrive in the more acidic oxygen minimum zone waters. On the other hand, evidence indicates that microbial anaerobic degradation of toluene may be a source of NO2− by partial denitrification, as shown for aquifer sediments. Since toluene production was not detected in incubations under aerobic conditions, we hypothesize that oxygen minimum zone waters export toluene to surrounding oxygenated waters. Expansion of hypoxia in the ocean will certainly enhance the production and export of anaerobic metabolites by marine microorganisms.