Substantial oxygen consumption by aerobic nitrite oxidation in oceanic oxygen minimum zones

Oceanic oxygen minimum zones (OMZs) are globally significant sites of biogeochemical cycling where microorganisms deplete dissolved oxygen (DO) to concentrations <20 µM. Amid intense competition for DO in these metabolically challenging environments, aerobic nitrite oxidation may consume significant amounts of DO and help maintain low DO concentrations, but this remains unquantified. Using parallel measurements of oxygen consumption rates and 15N-nitrite oxidation rates applied to both water column profiles and oxygen manipulation experiments, we show that the contribution of nitrite oxidation to overall DO consumption systematically increases as DO declines below 2 µM. Nitrite oxidation can account for all DO consumption only under DO concentrations <393 nM found in and below the secondary chlorophyll maximum. These patterns are consistent across sampling stations and experiments, reflecting coupling between nitrate reduction and nitrite-oxidizing Nitrospina with high oxygen affinity (based on isotopic and omic data). Collectively our results demonstrate that nitrite oxidation plays a pivotal role in the maintenance and biogeochemical dynamics of OMZs.

Major processes of the dissolved cobalt cycle in the North and equatorial Pacific Ocean

Over the past decade, the GEOTRACES and wider trace metal geochemical community have made substantial contributions towards constraining the marine cobalt (Co) cycle and its major biogeochemical processes. However, few Co speciation studies have been conducted in the North and equatorial Pacific Ocean, a vast portion of the world’s oceans by volume and an important endmember of deep thermohaline circulation. Dissolved Co (dCo) samples, including total dissolved and labile Co, were measured at-sea during the GEOTRACES Pacific meridional transect (GP15) along the 152° W longitudinal from 56° N to 20° S. Along this transect, upper ocean dCo was linearly correlated to dissolved phosphate (slope = 82 ± 2 µM:M) due to phytoplankton uptake and remineralization. As depth increased, dCo concentrations became increasingly decoupled from phosphate concentrations due to co-scavenging with manganese oxide particles in the mesopelagic. The transect revealed an organically-bound coastal source of dCo to the Alaskan Stream associated with low salinity waters. An intermediate-depth hydrothermal flux of dCo was observed off the Hawaiian coast at the Loihi Seamount, and the elevated dCo was correlated to estimated xs3He at and above the vent site; however, the Loihi Seamount likely did not represent a major source of Co to the Pacific basin. Elevated concentrations of dCo within oxygen minimum zones (OMZs) in the equatorial North and South Pacific were consistent with the suppression of oxidative scavenging, and we estimate that future deoxygenation could increase the OMZ dCo inventory by 13–28 % over the next century. In North Pacific Deep Water (NPDW), a fraction of elevated ligand-bound dCo appeared protected from scavenging by the high biogenic particle flux in the North Pacific basin. This finding is counter to previous expectations of low dCo concentrations in the deep Pacific due to scavenging over thermohaline circulation. Compared to a Co global biogeochemical model, the observed transect displayed more extreme inventories and fluxes of dCo than predicted by the model, suggesting a highly dynamic Pacific Co cycle.

Modeling the marine chromium cycle: new constraints on global-scale processes

Chromium (Cr) and its isotopes hold great promise as a tracer of past oxygenation and marine biological activity due to the contrasted chemical properties of its two main oxidation states, Cr(III) and Cr(VI), and the associated isotope fractionation during redox transformations. However, to date the marine Cr cycle remains poorly constrained due to insufficient knowledge about sources and sinks and the influence of biological activity on redox reactions. We therefore implemented the two oxidation states of Cr in the Bern3D Earth system model of intermediate complexity in order to gain an improved understanding on the mechanisms that modulate the spatial distribution of Cr in the ocean. Due to the computational efficiency of the Bern3D model we are able to explore and constrain the range of a wide array of parameters. Our model simulates vertical, meridional, and inter-basin Cr concentration gradients in good agreement with observations. We find a mean ocean residence time of Cr between 5 and 8 kyr and a benthic flux, emanating from sediment surfaces, of 0.1–0.2 nmol cm−2 yr−1, both in the range of previous estimates. We further explore the origin of regional model–data mismatches through a number of sensitivity experiments. These indicate that the benthic Cr flux may be substantially lower in the Arctic than elsewhere. In addition, we find that a refined representation of oxygen minimum zones and their potential to reduce Cr yield Cr(III) concentrations and Cr removal rates in these regions in much improved agreement with observational data. Yet, further research is required to better understand the processes that govern these critical regions for Cr cycling.

Organic Matter Supply and Utilization in Oxygen Minimum Zones

Organic matter (OM) plays a significant role in the formation of oxygen minimum zones (OMZs) and associated biogeochemical cycling. OM supply processes to the OMZ include physical transport, particle formation, and sinking as well as active transport by migrating zooplankton and nekton. In addition to the availability of oxygen and other electron acceptors, the remineralization rate of OM is controlled by its biochemical quality. Enhanced microbial respiration of OM can induce anoxic microzones in an otherwise oxygenated water column. Reduced OM degradation under low-oxygen conditions, on the other hand, may increase the CO2 storage time in the ocean. Understanding the interdependencies between OM and oxygen cycling is of high relevance for an ocean facing deoxygenation as a consequence of global warming. In this review, we describe OM fluxes into and cycling within two large OMZs associated with eastern boundary upwelling systems that differ greatly in the extent of oxygen loss: the highly oxygen-depleted OMZ in the tropical South Pacific and the moderately hypoxic OMZ in the tropical North Atlantic. We summarize new findings from a large German collaborative research project, Collaborative Research Center 754 (SFB 754), and identify knowledge gaps and future research priorities. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

The Physiology and Biogeochemistry of SUP05

The SUP05 clade of gammaproteobacteria (Thioglobaceae) comprises both primary producers and primary consumers of organic carbon in the oceans. Host-associated autotrophs are a principal source of carbon and other nutrients for deep-sea eukaryotes at hydrothermal vents, and their free-living relatives are a primary source of organic matter in seawater at vents and in marine oxygen minimum zones. Similar to other abundant marine heterotrophs, such as SAR11 and Roseobacter, heterotrophic Thioglobaceae use the dilute pool of osmolytes produced by phytoplankton for growth, including methylated amines and sulfonates. Heterotrophic members are common throughout the ocean, and autotrophic members are abundant at hydrothermal vents and in anoxic waters; combined, they can account for more than 50% of the total bacterial community. Studies of both cultured and uncultured representatives from this diverse family are providing novel insights into the shifting biogeochemical roles of autotrophic and heterotrophic bacteria that cross oxic–anoxic boundary layers in the ocean. Expected final online publication date for the Annual Review of Marine Science, Volume 14 is January 2022. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

Novel metagenome-assembled genomes involved in the nitrogen cycle from a Pacific oxygen minimum zone

Oxygen minimum zones (OMZs) are unique marine regions where broad redox gradients stimulate biogeochemical cycles. Despite the important and unique role of OMZ microbes in these cycles, they are less characterized than microbes from the oxic ocean. Here we recovered 39 high- and medium-quality metagenome-assembled genomes (MAGs) from the Eastern Tropical South Pacific OMZ. More than half of these MAGs were not represented at the species level among 2631 MAGs from global marine datasets. OMZ MAGs were dominated by denitrifiers catalyzing nitrogen loss and especially MAGs with partial denitrification metabolism. A novel bacterial genome with nitrate-reducing potential could only be assigned to the phylum level. A Marine-Group II archaeon was found to be a versatile denitrifier, with the potential capability to respire multiple nitrogen compounds including N2O. The newly discovered denitrifying MAGs will improve our understanding of microbial adaptation strategies and the evolution of denitrification in the tree of life.

Small sinking particles control anammox rates in the Peruvian oxygen minimum zone

Anaerobic oxidation of ammonium (anammox) in oxygen minimum zones (OMZs) is a major pathway of oceanic nitrogen loss. Ammonium released from sinking particles has been suggested to fuel this process. During cruises to the Peruvian OMZ in April–June 2017 we found that anammox rates are strongly correlated with the volume of small particles (128–512 µm), even though anammox bacteria were not directly associated with particles. This suggests that the relationship between anammox rates and particles is related to the ammonium released from particles by remineralization. To investigate this, ammonium release from particles was modelled and theoretical encounters of free-living anammox bacteria with ammonium in the particle boundary layer were calculated. These results indicated that small sinking particles could be responsible for ~75% of ammonium release in anoxic waters and that free-living anammox bacteria frequently encounter ammonium in the vicinity of smaller particles. This indicates a so far underestimated role of abundant, slow-sinking small particles in controlling oceanic nutrient budgets, and furthermore implies that observations of the volume of small particles could be used to estimate N-loss across large areas.

Modeling the marine chromium cycle: New constraints on global-scale processes

Chromium (Cr) and its isotopes hold great promise as tracer of past oxygenation and marine biological activity due to the different chemical properties of its two main oxidation states, Cr(III) and Cr(VI), and the associated fractionation during redox transformations. However, to date the Cr cycle remains poorly constrained due to insufficient knowledge about sources and sinks and the influence of biological activity on redox reactions. We therefore implemented the two oxidation states of Cr in the Bern3D Earth system model of intermediate complexity, in order to gain an improved understanding on the mechanisms that modulate the spatial distribution of Cr in the ocean. Due to the computational efficiency of the Bern3D model we are able to explore and constrain the range of a wide array of parameters. Our model simulates vertical, meridional, and inter-basin Cr concentration gradients in good agreement with observations. We find a mean ocean residence time of Cr between 5 and 8 kyr, and a benthic flux, emanating from all sediment surfaces, of of 0.1–0.2 nmol cm-2 yr-1, both in the range of previous estimates. We further explore the origin of regional model-data mismatches through a number of sensitivity experiments. These indicate that the benthic Cr flux may be substantially lower in the Arctic than elsewhere. In addition, we find that a refined representation of oxygen minimum zones and their Cr redox potential yields Cr(III) concentrations and Cr removal rates in much improved agreement with observational data.