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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. Biological carbon pump pathways combine to transport organic carbon into the deep ocean. This study shows that sinking particles sequester 4 Pg C, active transport sequesters 1 Pg C, and subduction sequesters 0.8 Pg C in the California Current Ecosystem.

Passive sinking of particulate organic matter (POM) is the main mechanism through which the biological pump transports surface primary production to the ocean interior. However, the contribution and variability of different biological sources to vertical export is not fully understood. Here, we use DNA metabarcoding of the 18S rRNA gene and particle interceptor traps (PITs) to characterize the taxonomic composition of particles sinking out of the photic layer in the California Current Ecosystem (CCE), a productive system with high export potential. The PITs included formalin-fixed and ‘live’ traps to investigate eukaryotic communities involved in the export and remineralization of sinking particles. Sequences affiliated with Radiolaria dominated the eukaryotic assemblage in fixed traps (90%), with Dinophyta and Metazoa making minor contributions. The prominence of Radiolaria decreased drastically in live traps, possibly due to selective consumption by copepods, heterotrophic nanoflagellates, and phaeodarians that were heavily enriched in these traps. These patterns were consistent across the water masses surveyed extending from the coast to offshore, despite major differences in productivity and trophic st

Enhanced vertical carbon transport (gravitational sinking and subduction) at mesoscale ocean fronts may explain the demonstrated imbalance of new production and sinking particle export in coastal upwelling ecosystems. Based on flux assessments from 238U:234Th disequilibrium and sediment traps, we found 2 to 3 times higher rates of gravitational particle export near a deep-water front (305 mg C·m−2·d−1) compared with adjacent water or to mean (nonfrontal) regional conditions. Elevated particle flux at the front was mechanistically linked to Fe-stressed diatoms and high mesozooplankton fecal pellet production. Using a data assimilative regional ocean model fit to measured conditions, we estimate that an additional ∼225 mg C·m−2·d−1 was exported as subduction of particle-rich water at the front, highlighting a transport mechanism that is not captured by sediment traps and is poorly quantified by most models and in situ measurements. Mesoscale fronts may be responsible for over a quarter of total organic carbon sequestration in the California Current and other coastal upwelling ecosystems.

The Southern Ocean contributes substantially to the global biological carbon pump (BCP). Salps in the Southern Ocean, in particular Salpa thompsoni , are important grazers that produce large, fast-sinking fecal pellets. Here, we quantify the salp bloom impacts on microbial dynamics and the BCP, by contrasting locations differing in salp bloom presence/absence. Salp blooms coincide with phytoplankton dominated by diatoms or prymnesiophytes, depending on water mass characteristics. Their grazing is comparable to microzooplankton during their early bloom, resulting in a decrease of ~1/3 of primary production, and negative phytoplankton rates of change are associated with all salp locations. Particle export in salp waters is always higher, ranging 2- to 8- fold (average 5-fold), compared to non-salp locations, exporting up to 46% of primary production out of the euphotic zone. BCP efficiency increases from 5 to 28% in salp areas, which is among the highest recorded in the global ocean. Gelatinous bloom-forming zooplankton—salps—alter microbial communities and quintuple the flux of sinking particles from the surface to the deep, strongly enhancing the ability of the ocean to sequester CO 2 .

In contrast to its productive coastal margins, the open-ocean Gulf of Mexico (GoM) is notable for highly stratified surface waters with extremely low nutrient and chlorophyll concentrations. Field campaigns in 2017 and 2018 identified low rates of turbulent mixing, which combined with oligotrophic nutrient conditions, give very low estimates for diffusive flux of nitrate into the euphotic zone (< 1 µmol N m −2 d −1 ). Estimates of local N 2 -fixation are similarly low. In comparison, measured export rates of sinking particulate organic nitrogen (PON) from the euphotic zone are 2 – 3 orders of magnitude higher (i.e. 462 – 1144 µmol N m −2 d −1 ). We reconcile these disparate findings with regional scale dynamics inferred independently from remote-sensing products and a regional biogeochemical model and find that laterally-sourced organic matter is sufficient to support >90% of open-ocean nitrogen export in the GoM. Results show that lateral transport needs to be closely considered in studies of biogeochemical balances, particularly for basins enclosed by productive coasts. The middle of the Gulf of Mexico is stratified and highly oligotrophic, yet there are anomalously high fluxes of sinking particulate matter from the euphotic zone. Here the authors show that lateral advection of organic matter supports nitrogen export in the Gulf of Mexico’s open ocean.

Observations of phytoplankton abundances and community structure are critical towards understanding marine ecosystems. Common approaches to determine group-specific abundances include measuring phytoplankton pigments with high-performance liquid chromatography and DNA-based metabarcoding. Increasingly, mRNA abundances with metatranscriptomics are also employed. As phytoplankton pigments are used to develop and validate remote sensing algorithms, further comparisons between pigments and other metrics are needed to validate the extent to which these measurements agree for group-specific abundances; however, most previous comparisons have been hindered by metabarcoding and metatranscriptomics solely producing relative abundance data. By employing quantitative approaches that express both 18S rRNA genes (DNA) and total mRNA as concentrations, we show that these measurements are related for several eukaryotic phytoplankton groups. We further propose that integration of these can be used to examine ecological patterns more deeply. For example, productivity-diversity relationships of both the whole community and individual groups show a dinoflagellate-driven negative trend rather than the commonly found unimodal pattern. Pigments are also shown to relate to certain harmful algal bloom-forming taxa as well as the expression of sets of genes. Altogether, these results suggest that potential models of pigment concentrations via hyperspectral remote sensing may enable improved assessments of global phytoplankton community structure. These assessments may further support the detection of harmful algal blooms and the development of Earth system models.

Zooplankton play an important role in global biogeochemistry, and their secondary production supports valuable fisheries of the world's oceans. Currently, zooplankton standing stocks cannot be estimated using remote sensing techniques. Hence, coupled physical–biogeochemical models (PBMs) provide an important tool for studying zooplankton on regional and global scales. However, evaluating the accuracy of zooplankton biomass estimates from PBMs has been a major challenge due to sparse observations. In this study, we configure a PBM for the Gulf of Mexico (GoM) from 1993 to 2012 and validate the model against an extensive combination of biomass and rate measurements. Spatial variability in a multidecadal database of mesozooplankton biomass for the northern GoM is well resolved by the model with a statistically significant (p < 0.01) correlation of 0.90. Mesozooplankton secondary production for the region averaged 66±8×109 kg C yr−1, equivalent to ∼10 % of net primary production (NPP), and ranged from 51 to 82×109 kg C yr−1, with higher secondary production inside cyclonic eddies and substantially reduced secondary production in anticyclonic eddies. Model results from the shelf regions suggest that herbivory is the dominant feeding mode for small mesozooplankton (< 1 mm), whereas larger mesozooplankton are primarily carnivorous. In open-ocean oligotrophic waters, however, both mesozooplankton groups show proportionally greater reliance on heterotrophic protists as a food source. This highlights an important role of microbial and protistan food webs in sustaining mesozooplankton biomass in the GoM, which serves as the primary food source for early life stages of many commercially important fish species, including tuna.

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) for 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 supply 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 consumption loses 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 demands, 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 should be revisited.

Gelatinous filter feeders (e.g., salps, doliolids, and pyrosomes) have high filtration rates and can feed at predator:prey size ratios exceeding 10,000:1, yet are seldom included in ecosystem or climate models. We investigated foodweb and trophic dynamics in the presence and absence of salp blooms using traditional productivity and grazing measurements combined with compound-specific isotopic analysis of amino acids estimation of trophic position during Lagrangian framework experiments in the Southern Ocean. Trophic positions of salps ranging 10–132 mm in size were 2.2 ± 0.3 (mean ± std) compared to 2.6 ± 0.4 for smaller (mostly crustacean) mesozooplankton. The mostly herbivorous salp trophic position was maintained despite biomass dominance of ~10-µm-sized primary producers. We show that potential energy flux to >10-cm organisms increases by approximately an order of magnitude when salps are abundant, even without substantial alteration to primary production. Comparison to a wider dataset from other marine regions shows that alterations to herbivore communities are a better predictor of ecosystem transfer efficiency than primary-producer dynamics. These results suggest that diverse consumer communities and intraguild predation complicate climate change predictions (e.g., trophic amplification) based on linear food chains. These compensatory foodweb dynamics should be included in models that forecast marine ecosystem responses to warming and reduced nutrient supply. Food-web and isotope-based trophic analyses suggest that blooms of salps (gelatinous grazers) can increase ecosystem efficiency and energy transfer to large organisms. These compensatory food-web dynamics may offset predicted trophic amplification.

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.