Quantifying biological carbon pump pathways with a data-constrained mechanistic model ensemble approach

The ability to constrain the mechanisms that transport organic carbon into the deep ocean is complicated by the multiple physical, chemical, and ecological processes that intersect to create, transform, and transport particles in the ocean. In this manuscript we develop and parameterize a data-assimilative model of the multiple pathways of the biological carbon pump (NEMUROBCP). The mechanistic model is designed to represent sinking particle flux, active transport by vertically migrating zooplankton, and passive transport by subduction and vertical mixing, while also explicitly representing multiple biological and chemical properties measured directly in the field (including nutrients, phytoplankton and zooplankton taxa, carbon dioxide and oxygen, nitrogen isotopes, and 234Thorium). Using 30 different data types (including standing stock and rate measurements related to nutrients, phytoplankton, zooplankton, and non-living organic matter) from Lagrangian experiments conducted on 11 cruises from four ocean regions, we conduct an objective statistical parameterization of the model and generate one million different potential parameter sets that are used for ensemble model simulations. The model simulates in situ parameters that were assimilated (net primary production and gravitational particle flux) and parameters that were withheld (234Thorium and nitrogen isotopes) with reasonable accuracy. Model results show that gravitational flux of sinking particles and vertical mixing of organic matter from the surface ocean are more important biological pump pathways than active transport by vertically-migrating zooplankton. However, these processes are regionally variable, with sinking particles most important in oligotrophic areas of the Gulf of Mexico and California, sinking particles and vertical mixing roughly equivalent in productive regions of the CCE and the subtropical front in the Southern Ocean, and active transport an important contributor in the Eastern Tropical Pacific. We further find that mortality at depth is an important component of active transport when mesozooplankton biomasses are high, but that it is negligible in regions with low mesozooplankton biomass. Our results also highlight the high degree of uncertainty, particularly amongst mesozooplankton functional groups, that is derived from uncertainty in model parameters, with important implications from results that rely on non-ensemble model outputs. We also discuss the implications of our results for other data assimilation approaches.

210Po-210Pb Disequilibrium in the Western North Pacific Ocean: Particle Cycling and POC Export

Estimating the particulate organic carbon (POC) export flux from the upper ocean is fundamental for understanding the efficiency of the biological carbon pump driven by sinking particles in the oceans. The downward POC flux from the surface ocean based on 210Po-210Pb disequilibria in seawater samples from the western North Pacific Ocean (w-NPO) was measured in the early summer (May-June) of 2018. All the profiles showed a large 210Po deficiency relative to 210Pb in the euphotic zone (0–150 m), while this 210Po deficiency vanished below ∼500 m (with 210Po/210Pb ∼1 or > 1). A one-dimensional steady-state irreversible scavenging model was used to quantify the scavenging and removal fluxes of 210Po and 210Pb in the euphotic zone of the w-NPO. In the upper ocean (0–150 m), dissolved 210Po (D-Po) was scavenged into particles with a residence time of 0.6–5.5 year, and the 210Po export flux out of the euphotic zone was estimated as (0.33–3.49) × 104 dpm/m2/year, resulting in a wide range of particulate 210Po (P-Po) residence times (83–921 days). However, in the deep ocean (150–1,000 m), 210Po was transferred from the particulate phase to the dissolved phase. Using an integrated POC inventory and the P-Po residence times (Eppley model) in the w-NPO euphotic zone, the POC export fluxes (mmol C/m2/d) varied from 0.6 ± 0.2 to 8.8 ± 0.4. In comparison, applying the POC/210Po ratio of all (>0.45 μm) particles to 210Po export flux (Buesseler model), the obtained POC export fluxes (mmol C/m2/d) ranged from 0.7 ± 0.1 to 8.6 ± 0.8. Both Buesseler and Eppley methods showed enhanced POC export fluxes at stations near the continental shelf (i.e., Luzon Strait and the Oyashio-Kuroshio mixing region). The Eppley model-based 210Po-derived POC fluxes agreed well with the Buesseler model-based fluxes, indicating that both models are suitable for assessing POC fluxes in the w-NPO. The POC export efficiency was < 15%, suggesting a moderate biological carbon pump efficiency in the w-NPO. These low export efficiencies may be associated with the dominance of smaller particles and the processes of degradation and subsequent remineralization of these small particles in the euphotic zone of oligotrophic regions in the w-NPO.

Sea-ice derived meltwater stratification slows the biological carbon pump: results from continuous observations

The ocean moderates the world’s climate through absorption of heat and carbon, but how much carbon the ocean will continue to absorb remains unknown. The North Atlantic Ocean west (Baffin Bay/Labrador Sea) and east (Fram Strait/Greenland Sea) of Greenland features the most intense absorption of anthropogenic carbon globally; the biological carbon pump (BCP) contributes substantially. As Arctic sea-ice melts, the BCP changes, impacting global climate and other critical ocean attributes (e.g. biodiversity). Full understanding requires year-round observations across a range of ice conditions. Here we present such observations: autonomously collected Eulerian continuous 24-month time-series in Fram Strait. We show that, compared to ice-unaffected conditions, sea-ice derived meltwater stratification slows the BCP by 4 months, a shift from an export to a retention system, with measurable impacts on benthic communities. This has implications for ecosystem dynamics in the future warmer Arctic where the seasonal ice zone is expected to expand.

Multi-year robotic observations reveal the seasonality of downward carbon export pathways in the Southern Ocean

At high latitudes, the export of organic matter from the surface to the ocean interior, the biological carbon pump, has conventionally been attributed to the gravitational sinking of particulate organic carbon (POC). Conspicuous deficits in ocean carbon budgets have recently challenged this long-lived paradigm of a sole pathway. Multiple strands of evidence have demonstrated the importance of additional export pathways, including the particle injection pumps (PIPs). Recent model estimates revealed that PIPs have a comparable downward POC flux to the biological gravitational pump (BGP), but with potentially different seasonal signatures. To date, logistical constraints have prevented concomitant and extensive observations of these pumps, and little is known about the seasonality of their fluxes. Here, using year-round robotic observations and recent advances in optical signal analysis, we concurrently investigated the functioning of two PIPs - the mixed layer and eddy subduction pumps - and the BGP in Southern Ocean waters. By comparing three phytoplankton bloom cycles in contrasting environments, we show how physical forcing and phytoplankton phenology influence the magnitude and seasonality of these pumps, with implications for carbon sequestration efficiency.

Decreasing Phanerozoic extinction intensity as a consequence of Earth surface oxygenation and metazoan ecophysiology

The decline in background extinction rates of marine animals through geologic time is an established but unexplained feature of the Phanerozoic fossil record. There is also growing consensus that the ocean and atmosphere did not become oxygenated to near-modern levels until the mid-Paleozoic, coinciding with the onset of generally lower extinction rates. Physiological theory provides us with a possible causal link between these two observations—predicting that the synergistic impacts of oxygen and temperature on aerobic respiration would have made marine animals more vulnerable to ocean warming events during periods of limited surface oxygenation. Here, we evaluate the hypothesis that changes in surface oxygenation exerted a first-order control on extinction rates through the Phanerozoic using a combined Earth system and ecophysiological modeling approach. We find that although continental configuration, the efficiency of the biological carbon pump in the ocean, and initial climate state all impact the magnitude of modeled biodiversity loss across simulated warming events, atmospheric oxygen is the dominant predictor of extinction vulnerability, with metabolic habitat viability and global ecophysiotype extinction exhibiting inflection points around 40% of present atmospheric oxygen. Given this is the broad upper limit for estimates of early Paleozoic oxygen levels, our results are consistent with the relative frequency of high-magnitude extinction events (particularly those not included in the canonical big five mass extinctions) early in the Phanerozoic being a direct consequence of limited early Paleozoic oxygenation and temperature-dependent hypoxia responses.

A largely invariant marine dissolved organic carbon reservoir across Earth's history

Marine dissolved organic carbon (DOC), the largest pool of reduced carbon in the oceans, plays an important role in the global carbon cycle and contributes to the regulation of atmospheric oxygen and carbon dioxide abundances. Despite its importance in global biogeochemical cycles, the long-term history of the marine DOC reservoir is poorly constrained. Nonetheless, significant changes to the size of the oceanic DOC reservoir through Earth’s history have been commonly invoked to explain changes to ocean chemistry, carbon cycling, and marine ecology. Here, we present a revised view of the evolution of marine DOC concentrations using a mechanistic carbon cycle model that can reproduce DOC concentrations in both oxic and anoxic modern environments. We use this model to demonstrate that the overall size of the marine DOC reservoir has likely undergone very little variation through Earth’s history, despite major changes in the redox state of the ocean–atmosphere system and the nature and efficiency of the biological carbon pump. A relatively static marine DOC reservoir across Earth’s history renders it unlikely that major changes in marine DOC concentrations have been responsible for driving massive repartitioning of surface carbon or the large carbon isotope excursions observed in Earth’s stratigraphic record and casts doubt on previously hypothesized links between marine DOC levels and the emergence and radiation of early animals.

Ecosystem function after the K/Pg extinction: decoupling of marine carbon pump and diversity

The ocean biological pump is the mechanism by which carbon and nutrients are transported to depth. As such, the biological pump is critical in the partitioning of carbon dioxide between the ocean and atmosphere, and the rate at which that carbon can be sequestered through burial in marine sediments. How the structure and function of planktic ecosystems in the ocean govern the strength and efficiency of the biological pump and its resilience to disruption are poorly understood. The aftermath of the impact at the Cretaceous/Palaeogene (K/Pg) boundary provides an ideal opportunity to address these questions as both the biological pump and marine plankton size and diversity were fundamentally disrupted. The excellent fossil record of planktic foraminifera as indicators of pelagic-biotic recovery combined with carbon isotope records tracing biological pump behaviour, show that the recovery of ecological traits (diversity, size and photosymbiosis) occurred much later (approx. 4.3 Ma) than biological pump recovery (approx. 1.8 Ma). We interpret this decoupling of diversity and the biological pump as an indication that ecosystem function had sufficiently recovered to drive an effective biological pump, at least regionally in the South Atlantic.