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Multiple processes transport carbon into the deep ocean as part of the biological carbon pump, leading to long-term carbon sequestration. However, our ability to predict future changes in these processes is hampered by the absence of studies that have simultaneously quantified all carbon pump pathways. Here, we quantify carbon export and sequestration in the California Current Ecosystem resulting from (1) sinking particles, (2) active transport by diel vertical migration, and (3) the physical pump (subduction + vertical mixing of particles). We find that sinking particles are the most important and export 9.0 mmol C m-2 d-1 across 100-m depth while sequestering 3.9 Pg C. The physical pump exports more carbon from the shallow ocean than active transport (3.8 vs. 2.9 mmol C m-2 d-1), although active transport sequesters more carbon (1.0 vs. 0.8 Pg C) because of deeper remineralization depths. We discuss the implications of these results for understanding biological carbon pump responses to climate change.

We investigated variability in the C:234Th ratio of sinking particles and its relationship to changing water column characteristics and plankton ecological dynamics during 29 Lagrangian experiments conducted on six cruises of the California Current Ecosystem Long-Term Ecological Research (CCE-LTER) Program. C:234Th ratios of sinking particles collected by a surface-tethered sediment trap (C:234ThST) varied from 2.3 to 20.5 μmol C dpm−1 over a depth range of 47–150 m. C:234ThST was significantly greater (by a factor of 1.8) than C:234Th ratios of suspended >51-μm particles collected in the same water parcels with in situ pumps. C:234Th ratios of large (>200-μm) sinking particles also exceeded those of smaller sinking particles. C:234ThST decreased with depth from the base of the euphotic zone through the upper twilight zone. C:234ThST was positively correlated with several indices of ecosystem productivity including particulate organic carbon (POC) and chlorophyll (Chl) concentrations, mesozooplankton biomass, and the fraction of Chl >20-μm. Principal component analysis and multiple linear regression suggested that decaying phytoplankton blooms exhibited higher C:234ThST than actively growing blooms at similar biomass levels. C:234ThST was positively correlated with indices of the fractional contribution of fecal pellets in sediment traps when the proportion of fecal pellets was low in the traps, likely because of a correlation between mesozooplankton biomass and other indices of ecosystem productivity. However, when fecal pellets were a more important component of sinking material, C:234ThST decreased with increasing fecal pellet content. C:234ThST was also positively correlated with the Si:C ratio of sinking particles. Across the dataset (and across depths) a strong correlation was found between C:234ThST and the ratio of vertically-integrated POC to vertically-integrated total water column 234Th (vC:234Thtot). A mechanistic one-layer, two-box model of thorium sorption and desorption was invoked to explain this correlation. Two empirical models (one using vC:234Thtot; one using depth and vertically-integrated Chl) were developed to predict C:234Th ratios in this coastal upwelling biome. The former regression (log10(C:234ThST) = 0.43 × log10(vC:234Thtot) + 0.53) was found to also be a reasonable predictor for C:234ThST from diverse regions including the Southern Ocean, Sargasso Sea, Subarctic North Pacific, and Eastern Tropical North Pacific.

Zooplankton are important consumers of sinking particles in the ocean's twilight zone. However, the impact of different taxa depends on their feeding mode. In contrast to typical suspension-feeding zooplankton, flux-feeding taxa preferentially consume rapidly sinking particles that would otherwise penetrate into the deep ocean. To quantify the potential impact of two flux-feeding zooplankton taxa [Aulosphaeridae (Rhizaria), and Limacina helicina (euthecosome pteropod)] and the total suspension-feeding zooplankton community, we measured depth-stratified abundances of these organisms during six cruises in the California Current Ecosystem. Using allometric-scaling relationships, we computed the percentage of carbon flux intercepted by flux feeders and suspension feeders. These estimates were compared to direct measurements of carbon flux attenuation (CFA) made using drifting sediment traps and 238U-234Th disequilibrium. We found that CFA in the shallow twilight zone typically ranged from 500 to 1000 μmol organic C flux remineralized per 10-m vertical depth bin. This equated to approximately 6-10% of carbon flux remineralized/10 m. The two flux-feeding taxa considered in this study could account for a substantial proportion of this flux near the base of the euphotic zone. The mean flux attenuation attributable to Aulosphaeridae was 0.69%/10 m (median = 0.21%/10 m, interquartile range = 0.04-0.81%) at their depth of maximum abundance (~100 m), which would equate to ~10% of total flux attenuation in this depth range. The maximum flux attenuation attributable to Aulosphaeridae reached 4.2%/10 m when these protists were most abundant. L. helicina, meanwhile, could intercept 0.45-1.6% of carbon flux/10 m, which was slightly greater (on average) than the Aulosphaeridae. In contrast, suspension-feeding zooplankton in the mesopelagic (including copepods, euphausiids, appendicularians, and ostracods) had combined clearance rates of 2-81 L m-3 day-1 (mean of 19.6 L m-3 day-1). This implies a substantial impact on slowly sinking particles, but a negligible impact on the presumably rapidly sinking fecal pellets that comprised the majority of the material collected in sediment traps. Our results highlight the need for a greater research focus on the many taxa that potentially act as flux feeders in the oceanic twilight zone.

We used extensive ecological and biogeochemical measurements obtained from quasi-Lagrangian experiments during two California Current Ecosystem Long-Term Ecosystem Research cruises to analyze carbon fluxes between the epipelagic and mesopelagic zones using a linear inverse ecosystem model (LIEM). Measurement constraints on the model include 14C primary productivity, dilution-based microzooplankton grazing rates, gut pigment-based mesozooplankton grazing rates (on multiple zooplankton size classes), 234Th:238U disequilibrium and sediment trap measured carbon export, and metabolic requirements of micronekton, zooplankton, and bacteria. A likelihood approach (Markov Chain Monte Carlo) was used to estimate the resulting flow uncertainties from a sample of potential flux networks. Results highlight the importance of mesozooplankton active transport (i.e., diel vertical migration) in supplying the carbon demand of mesopelagic organisms and sequestering carbon dioxide from the atmosphere. In nine water parcels ranging from a coastal bloom to offshore oligotrophic conditions, mesozooplankton active transport accounted for 18–84% (median: 42%) of the total carbon transfer to the mesopelagic, with gravitational settling of POC (12–55%; median: 37%), and subduction (2–32%; median: 14%) providing the majority of the remainder. Vertically migrating zooplankton contributed to downward carbon flux through respiration and excretion at depth and via mortality losses to predatory zooplankton and mesopelagic fish (e.g., myctophids and gonostomatids). Sensitivity analyses showed that the results of the LIEM were robust to changes in nekton metabolic demand, rates of bacterial production, and mesozooplankton gross growth efficiency. This analysis suggests that prior estimates of zooplankton active transport based on conservative estimates of standard (rather than active) metabolism are likely too low.

Sinking particles transport organic carbon produced in the surface ocean to the ocean interior, leading to net storage of atmospheric CO2 in the deep ocean. The rapid growth of in situ imaging technology has the potential to revolutionize our understanding of particle flux attenuation in the ocean; however, estimating particle flux from particle size and abundance (measured directly by in situ cameras) is challenging. Sinking rates are dependent on several factors, including particle excess density and porosity, which vary based on particle origin and type. Additionally, particle characteristics are transformed while sinking. We compare optically measured particle size spectra profiles (Underwater Vision Profiler 5, UVP) with contemporaneous measurements of particle flux made using sediment traps and 234Th:238U disequilibrium on six process cruises from the California Current Ecosystem (CCE) LTER Program. These measurements allow us to assess the efficacy of size-flux relationships for estimating fluxes from optical particle size measurements. We find that previously published parameterizations that estimate carbon flux from UVP profiles are a poor fit to direct flux measurements in the CCE. This discrepancy is found to result primarily from the important role of fecal pellets in particle flux. These pellets are primarily in a size range (i.e., 100–400 μm) that is not well-resolved as images by the UVP due to the resolution of the sensor. We develop new, CCE-optimized parameters for use in an algorithm estimating carbon flux from UVP data in the southern California Current (Flux = (Formula presented.)), with A = 15.4, B = 1.05, d = particle diameter (mm) and Flux in units of mg C m–2 d–1. We caution, however, that increased accuracy in flux estimates derived from optical instruments will require devices with greater resolution, the ability to differentiate fecal pellets from low porosity marine snow aggregates, and improved sampling of rapidly sinking fecal pellets. We also find that the particle size-flux relationships may be different within the euphotic zone than in the shallow twilight zone and hypothesize that the changing nature of sinking particles with depth must be considered when investigating the remineralization length scale of sinking particles in the ocean.

Predicting marine carbon sequestration in a changing climate requires mechanistic understanding of the processes controlling sinking particle flux under different climatic conditions. The recent occurrence of a warm anomaly (2014–2015) followed by an El Niño (2015–2016) in the southern sector of the California Current System presented an opportunity to analyze changes in the biological carbon pump in response to altered climate forcing. We compare primary production, mesozooplankton grazing, and carbon export from the euphotic zone during quasi-Lagrangian experiments conducted in contrasting conditions: two cruises during warm years - one during the warm anomaly in 2014 and one toward the end of El Niño 2016 – and three cruises during El Niño-neutral years. Results showed no substantial differences in the relationships between vertical carbon export and its presumed drivers (primary production, mesozooplankton grazing) between warm and neutral years. Mesozooplankton fecal pellet enumeration and phaeopigment measurements both showed that fecal pellets were the dominant contributor to export in productive upwelling regions. In more oligotrophic regions, fluxes were dominated by amorphous marine snow with negligible pigment content. We found no evidence for a significant shift in the relationship between mesozooplankton grazing rate and chlorophyll concentration. However, mass-specific grazing rates were lower at low-to-moderate chlorophyll concentrations during warm years relative to neutral years. We also detected a significant difference in the relationship between phytoplankton primary production and photosynthetically active radiation between years: at similar irradiance and nutrient concentrations, productivity decreased during the warm events. Whether these changes resulted from species composition changes remains to be determined. Overall, our results suggest that the processes driving export remain similar during different climate conditions, but that species compositional changes or other structural changes require further attention.

During two cruises in the oligotrophic oceanic Gulf of Mexico, we deployed sediment traps at three depths: center of the euphotic zone (EZ) (60 m), base of the EZ (117–151 m), and in the twilight zone (231 m). Organic carbon export declined with depth from 6.4 to 4.6 to 2.4 mmol C m−2 d−1, suggesting that net particle production was concentrated in the upper EZ. Net primary production varied from 24 to 29 mmol C m−2 d−1, slightly more than half in the upper EZ. Export ratios varied from 11 to 25%. Trap measurements of chlorophyll and phaeopigments allowed us to quantify fluxes of fresh phytoplankton and herbivorous fecal pellets, respectively, which were both minor contributors to total flux, although their contributions varied with depth. Phytoplankton flux was more important from the upper to lower EZ; fecal pellets were more important at the EZ base and below. C:N elemental ratios and 13C and 15N isotope analyses indicated particle transformations within and beneath the EZ. 234Th-238U disequilibrium measurements varied, likely reflecting the mixing of water from multiple regions over the ~month-long time-scale of 234Th. Our results highlight the complexity of the biological carbon pump in oligotrophic regions. [ABSTRACT FROM AUTHOR]

The highly stratified, oligotrophic regions of the oceans are predominantly nitrogen limited in the surface ocean and light limited at the deep chlorophyll maximum (DCM). Hence, determining light and nitrogen co-limitation patterns for diverse phytoplankton taxa is crucial to understanding marine primary production throughout the euphotic zone. During two cruises in the deep-water Gulf of Mexico, we measured primary productivity (H13CO3−), nitrate uptake (15NO3−) and ammonium uptake (15NH4+) throughout the water column. Primary productivity declined with depth from the mixed layer to the DCM, averaging 27.1 mmol C m−2 d−1. The fraction of growth supported by NO3− was consistently low, with upper euphotic zone values ranging from 0.01 to 0.14 and lower euphotic zone values ranging from 0.03 to 0.44. Nitrate uptake showed strong diel patterns (maximum during the day), whereas ammonium uptake exhibited no diel variability. To parameterize taxon-specific phytoplankton nutrient and light utilization, we used a data assimilation approach (Bayesian Markov Chain Monte Carlo) including primary productivity, nutrient uptake and taxon-specific growth rate measurements. Parameters derived from this analysis define distinct niches for five phytoplankton taxa (Prochlorococcus , Synechococcus , diatoms, dinoflagellates and prymnesiophytes) and may be useful for constraining biogeochemical models of oligotrophic open-ocean systems. [ABSTRACT FROM AUTHOR]

The availability of nitrogen (N) in ocean surface waters affects rates of photosynthesis and marine ecosystem structure. In spite of low dissolved inorganic N concentrations, export production in oligotrophic waters is comparable to more nutrient replete regions. Prior observations raise the possibility that di-nitrogen (N2) fixation supplies a significant fraction of N supporting export production in the Gulf of Mexico. In this study, geochemical tools were used to quantify the relative and absolute importance of both subsurface nitrate and N2 fixation as sources of new N fueling export production in the oligotrophic Gulf of Mexico in May 2017 and May 2018. Comparing the isotopic composition ("δ15N") of nitrate with the δ15N of sinking particulate N collected during five sediment trap deployments each lasting two to four days indicates that N2 fixation is typically not detected and that the majority (≥80%) of export production is supported by subsurface nitrate. Moreover, no gradients in upper ocean dissolved organic N and suspended particulate N concentration and/or δ15N were found that would indicate significant N2 fixation fluxes accumulated in these pools, consistent with low Trichodesmium spp. abundance. Finally, comparing the δ15N of sinking particulate N captured within vs. below the euphotic zone indicates that during late spring regenerated N is low in δ15N compared to sinking N. [ABSTRACT FROM AUTHOR]

We used linear inverse ecosystem modeling techniques to assimilate data from extensive Lagrangian field experiments into a mass-balance constrained food web for the Gulf of Mexico open-ocean ecosystem. This region is highly oligotrophic, yet Atlantic bluefin tuna (ABT) travel long distances from feeding grounds in the North Atlantic to spawn there. Our results show extensive nutrient regeneration fueling primary productivity (mostly by cyanobacteria and other picophytoplankton) in the upper euphotic zone. The food web is dominated by the microbial loop (>70% of net primary productivity is respired by heterotrophic bacteria and protists that feed on them). By contrast, herbivorous food web pathways from phytoplankton to metazoan zooplankton process <10% of the net primary production in the mixed layer. Nevertheless, ABT larvae feed preferentially on podonid cladocerans and other suspension-feeding zooplankton, which in turn derive much of their nutrition from nano- and micro-phytoplankton (mixotrophic flagellates, and to a lesser extent, diatoms). This allows ABT larvae to maintain a comparatively low trophic level (~4.2 for preflexion and postflexion larvae), which increases trophic transfer from phytoplankton to larval fish. [ABSTRACT FROM AUTHOR]

Phytoplankton growth and microzooplankton grazing rates were measured in repeated profiles of dilution experiments incubated in situ on a drift array in order to assess microbial production and food web characteristics in the oligotrophic bluefin tuna spawning habitat of the Gulf of Mexico (May peak spawning seasons, 2017–2018). Grazing often exceeded growth with the processes more balanced overall in the surface mixed layer, but biomass accumulated in the mid-euphotic zone. Community production estimates (260–500 mg C m−2 day−1) were low compared to similar open-ocean studies in the Pacific Ocean. Prochlorococcus was a consistent major contributor (113–204 mg C m−2 day−1) to productivity, while diatoms and dinoflagellates (2–10 and 4–13 mg C m−2 day−1, respectively) were consistently low. Prymnesiophytes, the most dynamic component (34–134 mg C m−2 day−1), co-dominated in 2017 experiments. Unexpected imbalances in grazing relative to production were observed for all picoplankton populations (Prochlorococcus , Synechococcus and heterotrophic bacteria), suggesting a trophic cascade in the absence of mesozooplankton predation on large microzooplankton. Study sites with abundant larval tuna had the shallowest deep chlorophyll maxima and significant net positive phytoplankton growth below the mixed layer. [ABSTRACT FROM AUTHOR]

Biomass and composition of the phytoplankton community were investigated in the deep-water Gulf of Mexico (GoM) at the edges of Loop Current anticyclonic eddies during May 2017 and May 2018. Using flow cytometry, high-performance liquid chromatography pigments and microscopy, we found euphotic zone integrated chlorophyll a of ~10 mg m−2 and autotrophic carbon ranging from 463 to 1268 mg m−2, dominated by picoplankton (<2 μm cells). Phytoplankton assemblages were similar to the mean composition at the Bermuda Atlantic Time-series Study site, but differed from the Hawaii Ocean Times-series site. GoM phytoplankton biomass was ~2-fold higher at the deep chlorophyll maximum (DCM) relative to the mixed layer (ML). Prochlorococcus and prymnesiophytes were the dominant taxa throughout the euphotic zone; however, other eukaryotic taxa had significant biomass in the DCM. Shallower DCMs were correlated with more prymnesiophytes and prasinophytes (Type 3) and reduced Prochlorococcus. These trends in ML and DCM taxonomic composition likely reflect relative nutrient supply—with ML populations relying on remineralized ammonium as a nitrogen source, and the taxonomically diverse DCM populations using more nitrate. These spatially separated phytoplankton communities represent different pathways for primary production, with a dominance of picoplankton in the ML and more nano- and microplankton at the DCM. [ABSTRACT FROM AUTHOR]

The ratio of dissolved oxygen to argon in seawater is frequently employed to estimate rates of net community production (NCP) in the oceanic mixed layer. The in situ O2/Ar‐based method accounts for many physical factors that influence oxygen concentrations, permitting isolation of the biological oxygen signal produced by the balance of photosynthesis and respiration. However, this technique traditionally relies upon several assumptions when calculating the mixed‐layer O2/Ar budget, most notably the absence of vertical fluxes of O2/Ar and the principle that the air‐sea gas exchange of biological oxygen closely approximates net productivity rates. Employing a Lagrangian study design and leveraging data outputs from a regional physical oceanographic model, we conducted in situ measurements of O2/Ar in the California Current Ecosystem in spring 2016 and summer 2017 to evaluate these assumptions within a "worst‐case" field environment. Quantifying vertical fluxes, incorporating nonsteady state changes in O2/Ar, and comparing NCP estimates evaluated over several day versus longer timescales, we find differences in NCP metrics calculated over different time intervals to be considerable, also observing significant potential effects from vertical fluxes, particularly advection. Additionally, we observe strong diel variability in O2/Ar and NCP rates at multiple stations. Our results reemphasize the importance of accounting for vertical fluxes when interpreting O2/Ar‐derived NCP data and the potentially large effect of nonsteady state conditions on NCP evaluated over shorter timescales. In addition, diel cycles in surface O2/Ar can also bias interpretation of NCP data based on local productivity and the time of day when measurements were made. Plain Language Summary: Marine microbes produce and consume oxygen as a product of a balance between photosynthesis and respiration, thereby causing changes in the concentration of oxygen in the surface ocean. Taking advantage of known relationships between the gas properties of oxygen and argon, researchers can isolate changes in biologically produced oxygen from changes that result from physical processes. This calculation relies on several simplifying assumptions. In this study, we test the importance of these assumptions by tracking changes in biological oxygen while following several water parcels over the course of 2–4 days. We find that accounting for vertical mixing processes and the rate of change in biological oxygen is important when estimating biological oxygen production rates. We also find that daily shifts in oxygen as photosynthesis rises through the day and falls to zero at night can influence measurements based on time of collection. Key Points: We employ Lagrangian observations of O2/Ar in the California Current to test assumptions inherent to the O2/Ar‐based NCP methodWe demonstrate the significance of vertical fluxes and diel patterns in O2/Ar that can bias NCP measurementsNonsteady state conditions and vertical fluxes strongly complicate efforts to evaluate O2/Ar‐based NCP on short timescales [ABSTRACT FROM AUTHOR]

A multimethod process-oriented investigation of diverse productivity measures in the California Current Ecosystem (CCE) Long-Term Ecological Research study region, a complex physical environment, is presented. Seven multiday deployments covering a transition region from high to low productivity were conducted over two field expeditions (spring 2016 and summer 2017). Employing a Lagrangian study design, water parcels were followed over several days, comparing 24-h in situ measurements (14C and 15NO3-uptake, dilution estimates of phytoplankton growth, and microzooplankton grazing) with high-resolution productivity measurements by fast repetition rate fluorometry (FRRF) and equilibrium inlet mass spectrometry (EIMS), and integrated carbon export measuremnts using sediment traps. Results show the importance of accounting for temporal and fine spatial scale variability when estimating ecosystem production. FRRF and EIMS measurements resolved diel patterns in gross primary and net community production. Diel productivity changes agreed well with comparably more traditional measurements. While differences in productivity metrics calculated over different time intervals were considerable, as those methods rely on different base assumptions, the data can be used to explain ecosystem processes which would otherwise have gone unnoticed. The processes resolved from this method comparison further understanding of temporal and spatial coupling and decoupling of surface productivity and potential carbon burial in a gradient from coastal to offshore ecosystems. [ABSTRACT FROM AUTHOR]

Oceanographic field programs often use δ15N biogeochemical measurements and in situ rate measurements to investigate nitrogen cycling and planktonic ecosystem structure. However, integrative modeling approaches capable of synthesizing these distinct measurement types are lacking. We develop a novel approach for incorporating δ15N isotopic data into existing Markov Chain Monte Carlo (MCMC) random walk methods for solving linear inverse ecosystem models. We test the ability of this approach to recover food web indices (nitrate uptake, nitrogen fixation, zooplankton trophic level, and secondary production) derived from forward models simulating the planktonic ecosystems of the California Current and Amazon River Plume. We show that the MCMC with δ15N approach typically does a better job of recovering ecosystem structure than the standard MCMC or L2 minimum norm (L2MN) approaches, and also outperforms an L2MN with δ15N approach. Furthermore, we find that the MCMC with δ15N approach is robust to the removal of input equations and hence is well suited to typical pelagic ecosystem studies for which the system is usually vastly under-constrained. Our approach is easily extendable for use with δ13C isotopic measurements or variable carbon:nitrogen stoichiometry. [ABSTRACT FROM AUTHOR]