Lotze HK, Tittensor DP, Bryndum-Buchholz A, Cheung WWL, Galbraith ED, Barange M, Barrier N, Bianchi D, Blanchard JL, Bopp L, et al. Global ensemble projections reveal trophicamplification of ocean biomass declineswith climate change. Proceedings of the National Academy of Sciences [Internet]. 2019. Publisher's VersionAbstract

While the physical dimensions of climate change are now routinely assessed through multimodel intercomparisons, projected impacts on the global ocean ecosystem generally rely on individual models with a specific set of assumptions. To address these single-model limitations, we present standardized ensemble projections from six global marine ecosystem models forced with two Earth system models and four emis- sion scenarios with and without fishing. We derive average biomass trends and associated uncertainties across the marine food web. Without fishing, mean global animal biomass decreased by 5% (±4% SD) under low emissions and 17% (±11% SD) under high emissions by 2100, with an average 5% decline for every 1 °C of warming. Projected biomass declines were primarily driven by increasing temperature and decreasing primary production, and were more pronounced at higher trophic levels, a process known as trophic amplification. Fishing did not substantially alter the effects of climate change. Considerable regional variation featured strong biomass increases at high latitudes and de- creases at middle to low latitudes, with good model agreement on the direction of change but variable magnitude. Uncertainties due to vari- ations in marine ecosystem and Earth system models were similar. Ensemble projections performed well compared with empirical data, emphasizing the benefits of multimodel inference to project future outcomes. Our results indicate that global ocean animal biomass con- sistently declines with climate change, and that these impacts are am- plified at higher trophic levels. Next steps for model development include dynamic scenarios of fishing, cumulative human impacts, and the effects of management measures on future ocean biomass trends.

Carozza DA, Bianchi D, Galbraith ED. Metabolic impacts of climate change on marine ecosystems: Implications for fish communities and fisheries. Global Ecology and Biogeography [Internet]. 2018.

Aim: Climate change will reshape marine ecosystems over the 21st century through diverse and complex mechanisms that are difficult to assess quantitatively. Here, we characterize expectations for how marine community biomass will respond to the energetic consequences of changes in primary production and temperature‐depend‐ ent metabolic rates, under a range of fishing/conservation scenarios.


Location: Global ocean.


Time period: 1950–2100.


Major taxa studied: Commercially harvested marine ectotherms (‘fish’).


Methods: We use a size‐structured macroecological model of the marine ecosystem, coupled with a catch model that allows for calibration with global historical data and simulation of fishing. We examine the four energetic mechanisms that, within the model framework, determine the community response to climate change: net pri‐ mary production, phytoplankton cell size, and the temperature dependencies of growth and natural mortality.


Results: Climate change decreases the modelled global fish community biomass by as much as 30% by 2100. This results from a diminished energy supply to upper trophic levels as photosynthesis becomes more nutrient limited and phytoplankton cells shrink, and from a temperature‐driven increase of natural mortality that, together, overwhelm the effect of accelerated somatic growth rates. Ocean circulation changes drive regional variations of primary production, producing patterns of winners and los‐ ers that largely compensate each other when averaged globally, whereas decreasing phytoplankton size drives weaker but more uniformly negative changes. The climate impacts are similar across the range of conservation scenarios but are slightly amplified in the strong conservation scenarios owing to the greater role of natural mortality. Main conclusions: The spatial pattern of climate impacts is mostly determined by changes in primary production. The overall decline of community biomass is attributed to a temperature‐driven increase of natural mortality, alongside an overall decrease in phytoplankton size, despite faster somatic growth. Our results highlight the importance of the competition between accelerated growth and mortality in a warming ocean.

Yamamoto A, Palter JB, Dufour CO, Griffies SM, Bianchi D, Claret M, Dunne JP, Frenger I, Galbraith ED. Roles of the ocean mesoscale in the horizontal supply of mass, heat, carbon and nutrients to the Northern Hemisphere subtropical gyres. Journal of Geophysical Research: Oceans [Internet]. 2018. Publisher's VersionAbstract

Horizontal transport at the boundaries of the subtropical gyres plays a crucial role in providing the nutrients that fuel gyre primary productivity, the heat that helps restratify the surface mixed layer, and the dissolved inorganic carbon that influences air‐sea carbon exchange. Mesoscale eddies may be an important component of these horizontal transports; however, previous studies have not quantified the horizontal tracer transport due to eddies across the subtropical gyre boundaries. Here we assess the physical mechanisms that control the horizontal transport of mass, heat, nutrients, and carbon across the North Pacific and North Atlantic subtropical gyre boundaries using the eddy‐rich ocean component of a climate model (Geophysical Fluid Dynamics Laboratory Climate Model version 2.6) coupled to a simple biogeochemical model (mini‐BLING). Our results suggest that horizontal transport across the gyre boundaries supplies a substantial amount of mass and tracers to the ventilated layer of both Northern Hemisphere subtropical gyres, with the Kuroshio and Gulf Stream acting as main exchange gateways. Mass, heat, and dissolved inorganic carbon supply is principally driven by the time‐mean circulation, whereas nutrient transport differs markedly from the other tracers, as nutrients are mainly supplied to both subtropical gyres by downgradient eddy mixing across gyre boundaries. A budget analysis further reveals that the horizontal nutrient transport, combining the roles of both mean and eddy components, is responsible for more than three quarters of the total nutrient supply into the subtropical gyres, surpassing a recent estimate based on a coarse‐resolution model and thus further highlighting the importance of horizontal nutrient transport.

Claret M, Galbraith ED, Palter JB, Bianchi D, Fennel K, Gilbert D, Dunne JP. Rapid coastal deoxygenation due to ocean circulation shift in the northwest Atlantic. Nature Climate Change [Internet]. 2018 :1. Publisher's VersionAbstract
Global observations show that the ocean lost approximately 2% of its oxygen inventory over the past five decades, with important implications for marine ecosystems. The rate of change varies regionally, with northwest Atlantic coastal waters showing a long-term drop that vastly outpaces the global and North Atlantic basin mean deoxygenation rates. However, past work has been unable to differentiate the role of large-scale climate forcing from that of local processes. Here, we use hydrographic evidence to show that a Labrador Current retreat is playing a key role in the deoxygenation on the northwest Atlantic shelf. A high-resolution global coupled climate–biogeochemistry model reproduces the observed decline of saturation oxygen concentrations in the region, driven by a retreat of the equatorward-flowing Labrador Current and an associated shift towards more oxygen-poor subtropical waters on the shelf. The dynamical changes underlying the shift in shelf water properties are correlated with a slowdown in the simulated Atlantic Meridional Overturning Circulation (AMOC). Our results provide strong evidence that a major, centennial-scale change of the Labrador Current is underway, and highlight the potential for ocean dynamics to impact coastal deoxygenation over the coming century.
Bianchi D, Weber TS, Kiko R, Deutsch C. Global niche of marine anaerobic metabolisms expanded by particle microenvironments. Nature Geoscience [Internet]. 2018. Publisher's VersionAbstract

In ocean waters, anaerobic microbial respiration should be confined to the anoxic waters found in coastal regions and tropical oxygen minimum zones, where it is energetically favourable. However, recent molecular and geochemical evidence has pointed to a much broader distribution of denitrifying and sulfate-reducing microbes. Anaerobic metabolisms are thought to thrive in microenvironments that develop inside sinking organic aggregates, but the global distribution and geochemical significance of these microenvironments is poorly understood. Here, we develop a new size-resolved particle model to predict anaerobic respiration from aggregate properties and seawater chemistry. Constrained by observations of the size spectrum of sinking particles, the model predicts that denitrification and sulfate reduction can be sustained throughout vast, hypoxic expanses of the ocean, and could explain the trace metal enrichment observed in particles due to sulfide precipitation. Globally, the expansion of the anaerobic niche due to particle microenvironments doubles the rate of water column denitrification compared with estimates based on anoxic zones alone, and changes the sensitivity of the marine nitrogen cycle to deoxygenation in a warming climate.

Frenger I, Bianchi D, Stührenberg C, Oschlies A, Dunne J, Deutsch C, Galbraith E, Schütte F. Biogeochemical role of subsurface coherent eddies in the ocean: Tracer cannonballs, hypoxic storms, and microbial stewpots?. Global Biogeochemical Cycles [Internet]. 2018;32 (2) :226-249. Publisher's VersionAbstract

Subsurface coherent eddies are well-known features of ocean circulation, but the sparsity of observations prevents an assessment of their importance for biogeochemistry. Here, we use a global eddying (0.1° ) ocean-biogeochemical model to carry out a census of subsurface coherent eddies originating from eastern boundary upwelling systems (EBUS), and quantify their biogeochemical effects as they propagate westward into the subtropical gyres. While most eddies exist for a few months, moving over distances of 100s of km, a small fraction (< 5%) of long-lived eddies propagates over distances greater than 1000km, carrying the oxygen-poor and nutrient-rich signature of EBUS into the gyre interiors. In the Pacific, transport by subsurface coherent eddies accounts for roughly 10% of the offshore transport of oxygen and nutrients in pycnocline waters. This "leakage" of subsurface waters can be a significant fraction of the transport by nutrient-rich poleward undercurrents, and may contribute to the well-known reduction of productivity by eddies in EBUS. Furthermore, at the density layer of their cores, eddies decrease climatological oxygen locally by close to 10%, thereby expanding oxygen minimum zones. Finally, eddies represent low-oxygen extreme events in otherwise oxygenated waters, increasing the area of hypoxic waters by several percent and producing dramatic short-term changes that may play an important ecological role. Capturing these non-local effects in global climate models, which typically include non-eddying oceans, would require dedicated parameterizations.

Galbraith ED, Carozza DA, Bianchi D. A coupled human-Earth model perspective on long-term trends in the global marine fishery. Nature Communications [Internet]. 2017;8 :14884. Publisher's VersionAbstract

The global wild marine fish harvest increased fourfold between 1950 and a peak value near the end of the 20th century, reflecting interactions between anthropogenic and ecological forces. Here, we examine these interactions in a bio-energetically constrained, spatially and temporally resolved model of global fisheries. We conduct historical hindcasts with the model, which suggest that technological progress can explain most of the 20th century increase of fish harvest. In contrast, projections extending this rate of technological progress into the future under open access suggest a long-term decrease in harvest due to over-fishing. Climate change is predicted to gradually decrease the global fish production capacity, though our model suggests that this is of secondary importance to social and economic factors. Our study represents a novel way to integrate human-ecological interactions within a single model framework for long-term simulations.

Carozza DA, Bianchi D, Galbraith ED. Formulation, General Features and Global Calibration of a Bioenergetically-Constrained Fishery Model. PloS one [Internet]. 2017;12 (1) :e0169763. Publisher's VersionAbstract

Human exploitation of marine resources is profoundly altering marine ecosystems, while climate change is expected to further impact commercially-harvested fish and other species. Although the global fishery is a highly complex system with many unpredictable aspects, the bioenergetic limits on fish production and the response of fishing effort to profit are both relatively tractable, and are sure to play important roles. Here we describe a generalized, coupled biological-economic model of the global marine fishery that represents both of these aspects in a unified framework, the BiOeconomic mArine Trophic Size-spectrum (BOATS) model. BOATS predicts fish production according to size spectra as a function of net primary production and temperature, and dynamically determines harvest spectra from the biomass density and interactive, prognostic fishing effort. Within this framework, the equilibrium fish biomass is determined by the economic forcings of catchability, ex-vessel price and cost per unit effort, while the peak harvest depends on the ecosystem parameters. Comparison of a large ensemble of idealized simulations with observational databases, focusing on historical biomass and peak harvests, allows us to narrow the range of several uncertain ecosystem parameters, rule out most parameter combinations, and select an optimal ensemble of model variants. Compared to the prior distributions, model variants with lower values of the mortality rate, trophic efficiency, and allometric constant agree better with observations. For most acceptable parameter combinations, natural mortality rates are more strongly affected by temperature than growth rates, suggesting different sensitivities of these processes to climate change. These results highlight the utility of adopting large-scale, aggregated data constraints to reduce model parameter uncertainties and to better predict the response of fisheries to human behaviour and climate change.

Holzer M, DeVries T, Bianchi D, Newton R, Schlosser P, Winckler G. Objective estimates of mantle 3He in the ocean and implications for constraining the deep ocean circulation. Earth and Planetary Science Letters [Internet]. 2017;458 :305 - 314. Publisher's VersionAbstract

Abstract Hydrothermal vents along the ocean's tectonic ridge systems inject superheated water and large amounts of dissolved metals that impact the deep ocean circulation and the oceanic cycling of trace metals. The hydrothermal fluid contains dissolved mantle helium that is enriched in 3He relative to the atmosphere, providing an isotopic tracer of the ocean's deep circulation and a marker of hydrothermal sources. This work investigates the potential for the 3He/4He isotope ratio to constrain the ocean's mantle 3He source and to provide constraints on the ocean's deep circulation. We use an ensemble of 11 data-assimilated steady-state ocean circulation models and a mantle helium source based on geographically varying sea-floor spreading rates. The global source distribution is partitioned into 6 regions, and the vertical profile and source amplitude of each region are varied independently to determine the optimal 3He source distribution that minimizes the mismatch between modeled and observed δ3He. In this way, we are able to fit the observed δ3He distribution to within a relative error of ∼15%, with a global 3He source that ranges from 640 to 850 mol yr−1, depending on circulation. The fit captures the vertical and interbasin gradients of the δ3He distribution very well and reproduces its jet-sheared saddle point in the deep equatorial Pacific. This demonstrates that the data-assimilated models have much greater fidelity to the deep ocean circulation than other coarse-resolution ocean models. Nonetheless, the modelled δ3He distributions still display some systematic biases, especially in the deep North Pacific where δ3He is overpredicted by our models, and in the southeastern tropical Pacific, where observed westward-spreading δ3He plumes are not well captured. Sources inferred by the data-assimilated transport with and without isopycnally aligned eddy diffusivity differ widely in the Southern Ocean, in spite of the ability to match the observed distributions of \CFCs\ and radiocarbon for either eddy parameterization.

Carozza DA, Bianchi D, Galbraith ED. The ecological module of BOATS-1.0: a bioenergetically constrained model of marine upper trophic levels suitable for studies of fisheries and ocean biogeochemistry. Geoscientific Model Development. 2016;9 (4) :1545–1565.Abstract

Environmental change and the exploitation of marine resources have had profound impacts on marine communities, with potential implications for ocean biogeochemistry and food security. In order to study such global-scale problems, it is helpful to have computationally efficient numerical models that predict the first-order features of fish biomass production as a function of the environment, based on empirical and mechanistic understandings of marine ecosystems. Here we describe the ecological module of the BiOeconomic mArine Trophic Size-spectrum (BOATS) model, which takes an Earth-system approach to modelling fish biomass at the global scale. The ecological model is designed to be used on an Earth-system model grid, and determines size spectra of fish biomass by explicitly resolving life history as a function of local temperature and net primary production. Biomass production is limited by the availability of photosynthetic energy to upper trophic levels, following empirical trophic efficiency scalings, and by well-established empirical temperature-dependent growth rates. Natural mortality is calculated using an empirical size-based relationship, while reproduction and recruitment depend on both the food availability to larvae from net primary production and the production of eggs by mature adult fish. We describe predicted biomass spectra and compare them to observations, and conduct a sensitivity study to determine how they change as a function of net primary production and temperature. The model relies on a limited number of parameters compared to similar modelling efforts, while retaining reasonably realistic representations of biological and ecological processes, and is computationally efficient, allowing extensive parameter-space analyses even when implemented globally. As such, it enables the exploration of the linkages between ocean biogeochemistry, climate, and upper trophic levels at the global scale, as well as a representation of fish biomass for idealized studies of fisheries.

Bianchi D, Mislan KAS. Global patterns of diel vertical migration times and velocities from acoustic data. Limnology and Oceanography [Internet]. 2016;61 (1) :353–364. Publisher's VersionAbstract

Diel vertical migrations (DVM) of zooplankton and micronekton are observed throughout the world ocean, where they influence ecological interactions and biogeochemical cycles. Despite their common occurrence, descriptions of the characteristics of these migrations are currently limited at the large scale. We analyze trajectories of migrations from a global dataset of acoustic backscatter to identify the large-scale patterns of the timing and speed of DVM. Sound scattering layers (SSL) leave the surface 21 ± 20 min before sunrise, and return to it 17 ± 23 min after sunset, while changes in bulk surface backscatter appear to be nearly synchronous to sunrise and sunset. Mean downward migrations (7.6 ± 3.6 cm s−1) are significantly faster than mean upward migrations (6.5 ± 3.5 cm s−1). Furthermore, coherent and predictable variations of migration properties at the scale of ocean basins are evident. These variations appear to be related to the depths of migration, such that deeper migrations, observed for example in the subtropical gyres, the western tropical Pacific and the Southern Ocean, show earlier departures and later arrivals than shallower migrations. Vertical velocities peak in the tropical and subtropical regions, and decline towards the poles, with the strongest declines observed in the North Pacific. Migration velocities are also correlated to migration depths, with deeper migrations being faster than shallow migrations. These new constraints on the characteristics of migrating SSL could help shed light on the physiological, ecological, and environmental controls that regulate DVM behavior.

    Cartapanis O, Bianchi D, Jaccard SL, Galbraith ED. Global pulses of organic carbon burial in deep-sea sediments during glacial maxima. Nature communications [Internet]. 2016;7. Publisher's VersionAbstract

    The burial of organic carbon in marine sediments removes carbon dioxide from the ocean–atmosphere pool, provides energy to the deep biosphere, and on geological timescales drives the oxygenation of the atmosphere. Here we quantify natural variations in the burial of organic carbon in deep-sea sediments over the last glacial cycle. Using a new data compilation of hundreds of sediment cores, we show that the accumulation rate of organic carbon in the deep sea was consistently higher (50%) during glacial maxima than during interglacials. The spatial pattern and temporal progression of the changes suggest that enhanced nutrient supply to parts of the surface ocean contributed to the glacial burial pulses, with likely additional contributions from more efficient transfer of organic matter to the deep sea and better preservation of organic matter due to reduced oxygen exposure. These results demonstrate a pronounced climate sensitivity for this global carbon cycle sink.

    Galbraith ED, Dunne JP, Gnanadesikan A, Slater RD, Sarmiento JL, Dufour CO, de Souza GF, Bianchi D, Claret M, Rodgers KB, et al. Complex functionality with minimal computation: Promise and pitfalls of reduced-tracer ocean biogeochemistry models. Journal of Advances in Modeling Earth Systems [Internet]. 2015;7 (4) :2012–2028. Publisher's VersionAbstract

    Earth System Models increasingly include ocean biogeochemistry models in order to predict changes in ocean carbon storage, hypoxia, and biological productivity under climate change. However, state-of-the-art ocean biogeochemical models include many advected tracers, that significantly increase the computational resources required, forcing a trade-off with spatial resolution. Here, we compare a state-of-the art model with 30 prognostic tracers (TOPAZ) with two reduced-tracer models, one with 6 tracers (BLING), and the other with 3 tracers (miniBLING). The reduced-tracer models employ parameterized, implicit biological functions, which nonetheless capture many of the most important processes resolved by TOPAZ. All three are embedded in the same coupled climate model. Despite the large difference in tracer number, the absence of tracers for living organic matter is shown to have a minimal impact on the transport of nutrient elements, and the three models produce similar mean annual preindustrial distributions of macronutrients, oxygen, and carbon. Significant differences do exist among the models, in particular the seasonal cycle of biomass and export production, but it does not appear that these are necessary consequences of the reduced tracer number. With increasing CO2, changes in dissolved oxygen and anthropogenic carbon uptake are very similar across the different models. Thus, while the reduced-tracer models do not explicitly resolve the diversity and internal dynamics of marine ecosystems, we demonstrate that such models are applicable to a broad suite of major biogeochemical concerns, including anthropogenic change. These results are very promising for the further development and application of reduced-tracer biogeochemical models that incorporate “sub-ecosystem-scale” parameterizations.

    Galbraith ED, Kwon EY, Bianchi D, Hain MP, Sarmiento JL. The impact of atmospheric pCO2 on carbon isotope ratios of the atmosphere and ocean. Global Biogeochemical Cycles [Internet]. 2015;29 (3) :307–324. Publisher's VersionAbstract

    It is well known that the equilibration timescale for the isotopic ratios 13C/12C and 14C/12C in the ocean mixed layer is on the order of a decade, 2 orders of magnitude slower than for oxygen. Less widely appreciated is the fact that the equilibration timescale is quite sensitive to the speciation of dissolved inorganic carbon (DIC) in the mixed layer, scaling linearly with the ratio DIC/CO2, which varies inversely with atmospheric pCO2. Although this effect is included in models that resolve the role of carbon speciation in air-sea exchange, its role is often unrecognized, and it is not commonly considered in the interpretation of carbon isotope observations. Here we use a global three-dimensional ocean model to estimate the redistribution of the carbon isotopic ratios between the atmosphere and ocean due solely to variations in atmospheric pCO2. Under Last Glacial Maximum (LGM) pCO2, atmospheric Δ14C is increased by ≈30‰ due to the speciation change, all else being equal, raising the surface reservoir age by about 250 years throughout most of the ocean. For 13C, enhanced surface disequilibrium under LGM pCO2 causes the upper ocean, atmosphere, and North Atlantic Deep Water δ13C to become at least 0.2‰ higher relative to deep waters ventilated by the Southern Ocean. Conversely, under high pCO2, rapid equilibration greatly decreases isotopic disequilibrium. As a result, during geological periods of high pCO2, vertical δ13C gradients may have been greatly weakened as a direct chemical consequence of the high pCO2, masquerading as very well ventilated or biologically dead Strangelove Oceans. The ongoing anthropogenic rise of pCO2 is accelerating the equilibration of the carbon isotopes in the ocean, lowering atmospheric Δ14C and weakening δ13C gradients within the ocean to a degree that is similar to the traditional fossil fuel “Suess” effect.

      Cabré A, Marinov I, Bernardello R, Bianchi D. Oxygen minimum zones in the tropical Pacific across CMIP5 models: mean state differences and climate change trends. Biogeosciences Discussions [Internet]. 2015;12 (8). Publisher's VersionAbstract

       We analyse simulations of the Pacific Ocean oxygen minimum zones (OMZs) from 11 Earth system model contributions to the Coupled Model Intercomparison Project Phase 5, focusing on the mean state and climate change projections. The simulations tend to overestimate the volume of the OMZs, especially in the tropics and Southern Hemisphere. Compared to observations, five models introduce incorrect meridional asymmetries in the distribution of oxygen including larger southern OMZ and weaker northern OMZ, due to interhemispheric biases in intermediate water mass ventilation. Seven models show too deep an extent of the tropical hypoxia compared to observations, stemming from a deficient equatorial ventilation in the upper ocean, combined with too large a biologically driven downward flux of particulate organic carbon at depth, caused by particle export from the euphotic layer that is too high and remineralization in the upper ocean that is too weak.  At interannual timescales, the dynamics of oxygen in the eastern tropical Pacific OMZ is dominated by biological consumption and linked to natural variability in the Walker circulation. However, under the climate change scenario RCP8.5, all simulations yield small and discrepant changes in oxygen concentration at mid depths in the tropical Pacific by the end of the 21st century due to an almost perfect compensation between warming-related decrease in oxygen saturation and decrease in biological oxygen utilization. Climate change projections are at odds with recent observations that show decreasing oxygen levels at mid depths in the tropical Pacific. Out of the OMZs, all the CMIP5 models predict a decrease of oxygen over most of the surface and deep ocean at low latitudes and over all depths at high latitudes due to an overall slow-down of ventilation and increased temperature.

      Babbin AR, Bianchi D, Jayakumar A, Ward BB. Rapid nitrous oxide cycling in the suboxic ocean. Science [Internet]. 2015;348 (6239) :1127–1129. Publisher's VersionAbstract

      Nitrous oxide (N2O) is a powerful greenhouse gas and a major cause of stratospheric ozone depletion, yet its sources and sinks remain poorly quantified in the oceans. We used isotope tracers to directly measure N2O reduction rates in the eastern tropical North Pacific. Because of incomplete denitrification, N2O cycling rates are an order of magnitude higher than predicted by current models in suboxic regions, and the spatial distribution suggests strong dependence on both organic carbon and dissolved oxygen concentrations. Furthermore, N2O turnover is 20 times higher than the net atmospheric efflux. The rapid rate of this cycling coupled to an expected expansion of suboxic ocean waters implies future increases in N2O emissions.

      Bianchi D, Babbin AR, Galbraith ED. Enhancement of anammox by the excretion of diel vertical migrators. Proceedings of the National Academy of Sciences [Internet]. 2014;111 (44) :15653–15658. Publisher's VersionAbstract

      Measurements show that anaerobic ammonium oxidation with nitrite (anammox) is a major pathway of fixed nitrogen removal in the anoxic zones of the open ocean. Anammox requires a source of ammonium, which under anoxic conditions could be supplied by the breakdown of sinking organic matter via heterotrophic denitrification. However, at many locations where anammox is measured, denitrification rates are small or undetectable. Alternative sources of ammonium have been proposed to explain this paradox, for example through dissimilatory reduction of nitrate to ammonium and transport from anoxic sediments. However, the relevance of these sources in open-ocean anoxic zones is debated. Here, we bring to attention an additional source of ammonium, namely, the daytime excretion by zooplankton and micronekton migrating from the surface to anoxic waters. We use a synthesis of acoustic data to show that, where anoxic waters occur within the water column, most migrators spend the daytime within them. Although migrators export only a small fraction of primary production from the surface, they focus excretion within a confined depth range of anoxic water where particle input is small. Using a simple biogeochemical model, we suggest that, at those depths, the source of ammonium from organisms undergoing diel vertical migrations could exceed the release from particle remineralization, enhancing in situ anammox rates. The contribution of this previously overlooked process, and the numerous uncertainties surrounding it, call for further efforts to evaluate the role of animals in oxygen minimum zone biogeochemistry.

      Galbraith ED, Kienast M, others. The acceleration of oceanic denitrification during deglacial warming. Nature Geoscience [Internet]. 2013;6 (7) :579–584. Publisher's VersionAbstract

      Over much of the ocean’s surface, productivity and growth are limited by a scarcity of bioavailable nitrogen. Sedimentary δ15N records spanning the last deglaciation suggest marked shifts in the nitrogen cycle during this time, but the quantification of these changes has been hindered by the complexity of nitrogen isotope cycling. Here we present a database of δ15N in sediments throughout the world’s oceans, including 2,329 modern seafloor samples, and 76 timeseries spanning the past 30,000 years. We show that the δ15N values of modern seafloor sediments are consistent with values predicted by our knowledge of nitrogen cycling in the water column. Despite many local deglacial changes, the globally averaged δ15N values of sinking organic matter were similar during the Last Glacial Maximum and Early Holocene. Considering the global isotopic mass balance, we explain these observations with the following deglacial history of nitrogen inventory processes. During the Last Glacial Maximum, the nitrogen cycle was near steady state. During the deglaciation, denitrification in the pelagic water column accelerated. The flooding of continental shelves subsequently increased denitrification at the seafloor, and denitrification reached near steady-state conditions again in the Early Holocene. We use a recent parameterization of seafloor denitrification to estimate a 30–120% increase in benthic denitrification between 15,000 and 8,000 years ago. Based on the similarity of globally averaged δ15N values during the Last Glacial Maximum and Early Holocene, we infer that pelagic denitrification must have increased by a similar amount between the two steady states.

      Gnanadesikan A, Bianchi D, Pradal M-A. Critical role for mesoscale eddy diffusion in supplying oxygen to hypoxic ocean waters. Geophysical Research Letters [Internet]. 2013;40 (19) :5194–5198. Publisher's VersionAbstract

      Estimates of the oceanic lateral eddy diffusion coefficient Aredi vary by more than an order of magnitude, ranging from less than a few hundred m2/s to thousands of m2/s. This uncertainty has first-order implications for the intensity of oceanic hypoxia, which is poorly simulated by the current generation of Earth System Models. Using satellite-based estimate of oxygen consumption in hypoxic waters to estimate the required diffusion coefficient for these waters gives a value of order 1000 m2/s. Varying Aredi across a suite of Earth System Models yields a broadly consistent result given a thermocline diapycnal diffusion coefficient of 1 × 10−5 m2/s.

      Bianchi D, Stock C, Galbraith ED, Sarmiento JL. Diel vertical migration: Ecological controls and impacts on the biological pump in a one-dimensional ocean model. Global Biogeochemical Cycles [Internet]. 2013;27 (2) :478–491. Publisher's VersionAbstract

      Diel vertical migration (DVM) of zooplankton and micronekton is widespread in the ocean and forms a fundamental component of the biological pump, but is generally overlooked in global models of the Earth system. We develop a parameterization of DVM in the ocean and integrate it with a size-structured NPZD model. We assess the model's ability to recreate ecosystem and DVM patterns at three well-observed Pacific sites, ALOHA, K2, and EQPAC, and use it to estimate the impact of DVM on marine ecosystems and biogeochemical dynamics. Our model includes the following: (1) a representation of migration dynamics in response to food availability and light intensity; (2) a representation of the digestive and metabolic processes that decouple zooplankton feeding from excretion, egestion, and respiration; and (3) a light-dependent parameterization of visual predation on zooplankton. The model captures the first-order patterns in plankton biomass and productivity across the biomes, including the biomass of migrating organisms. We estimate that realistic migratory populations sustain active fluxes to the mesopelagic zone equivalent to between 15% and 40% of the particle export and contribute up to half of the total respiration within the layers affected by migration. The localized active transport has important consequences for the cycling of oxygen, nutrients, and carbon. We highlight the importance of decoupling zooplankton feeding and respiration and excretion with depth for capturing the impact of migration on the redistribution of carbon and nutrients in the upper ocean.