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Subtropical oceans contribute significantly to global primary production, but the fate of the picophytoplankton that dominate in these low-nutrient regions is poorly understood. Working in the subtropical Mediterranean, we demonstrate that subduction of water at ocean fronts generates 3D intrusions with uncharacteristically high carbon, chlorophyll, and oxygen that extend below the sunlit photic zone into the dark ocean. These contain fresh picophytoplankton assemblages that resemble the photic-zone regions where the water originated. Intrusions propagate depth-dependent seasonal variations in microbial assemblages into the ocean interior. Strikingly, the intrusions included dominant biomass contributions from nonphotosynthetic bacteria and enrichment of enigmatic heterotrophic bacterial lineages. Thus, the intrusions not only deliver material that differs in composition and nutritional character from sinking detrital particles, but also drive shifts in bacterial community composition, organic matter processing, and interactions between surface and deep communities. Modeling efforts paired with global observations demonstrate that subduction can flux similar magnitudes of particulate organic carbon as sinking export, but is not accounted for in current export estimates and carbon cycle models. Intrusions formed by subduction are a particularly important mechanism for enhancing connectivity between surface and upper mesopelagic ecosystems in stratified subtropical ocean environments that are expanding due to the warming climate.

Mesopelagic fishes may contribute substantially to marine carbon export and sequestration. However, uncertainty in this contribution due to limited precision of mesopelagic biomass and bioenergetic rate estimates has not been thoroughly quantified for any study site. Datasets that can confront these challenges are rare, particularly for comparing fish-mediated carbon flux to other biological carbon pump pathways. Using data from a unique three-ship expedition in spring 2021 in the subarctic Northeast Atlantic Ocean, we compare carbon transported by adult fish, zooplankton, and sinking particles, and calculate uncertainty in the relative contribution of fishes. Results indicate biomass- and bioenergetic-based uncertainty contributed roughly equally to variance in estimated carbon transport. The plausible range of mesopelagic fish carbon flux spans an order of magnitude: 1.6–21 mg C m−2 d−1 to 200 m depth and 0.52–9.6 mg C m−2 d−1 to 500 m. Fishes contributed ∼0.52%–18% at 200 m to the total biological carbon pump, and ∼0.43%–13% at 500 m. Of the fish-mediated carbon transport to 200 m, ∼8%–30% is sequestered on climate-relevant time scales (>100 years). This reinforces that carbon transport should not be conflated with carbon sequestration. These findings have implications for prioritizing future empirical measurements, evaluating trade-offs in fisheries management, and understanding the role of fishes in the biological carbon pump. [ABSTRACT FROM AUTHOR]

Contributions to the biological pump that arise from the physical circulation are referred to as physical particle injection pumps. A synthesized view of how these physical pumps interact with each other and other components of the biological pump does not yet exist. Here, observations from a quasi‐Lagrangian float and an ocean glider, deployed in the Southern Ocean's Subantarctic Zone for one month during the spring bloom, offer insight into daily‐to‐monthly fluctuations in the mixed layer pump (MLP) and the eddy subduction pump (ESP). Estimated independently, each mechanism contributes intermittent export fluxes of roughly several hundred milligrams of particulate organic carbon (POC) per square meter per day. The glider‐based estimates indicate sustained weekly periods of MLP export fluxes across the base of the mixed layer with a magnitude of ∼450±110 ${\sim} 450\pm 110$ mg POC m−2 ${\mathrm{m}}^{-2}$ day−1 ${\text{day}}^{-1}$. Potential export fluxes from the ESP, based on a mixed layer instability scaling, occasionally exceed 400 mg POC m−2 ${\mathrm{m}}^{-2}$ day−1 ${\text{day}}^{-1}$, with export elevated due to both strong inferred vertical velocities and enhanced isopycnal slopes. Significant export fluxes from the ESP are localized to the edges of mesoscale eddies and to fronts, whereas the MLP acts more broadly due to the larger scales of atmospheric forcing. Regimes occur when MLP and ESP export fluxes can have either the same or opposite sign. Simple summation of fluxes from existing parameterizations of the two pumps likely misrepresents the total physical carbon flux. Insights into how mesoscale stirring and submesoscale velocities set POC vertical structure is a key target to reduce uncertainty in global carbon export fluxes. Plain Language Summary: The ocean influences the global carbon cycle by transferring carbon from the surface into the deep ocean, where it is sequestered from exchange with the atmosphere for periods of decades to millennia. Marine organisms enhance this downward transfer by fixing dissolved CO2 ${\text{CO}}_{2}$ into organic matter during photosynthesis (at the ocean surface) and removing the carbon through sinking of organic matter or via consumption and defecation by higher trophic levels. These processes are collectively referred to as the biological pump. Processes controlled by the ocean circulation, termed "physical" processes, can also influence the vertical transfer of fixed carbon to the ocean interior. Various proposed physical carbon pump mechanisms have been assessed independently without considering potential interactions between them. Here, we use data sets collected from different autonomous vehicles deployed in the Southern Ocean to provide a first look at interactions between two physical carbon pumps: the mixed layer and eddy subduction pumps. These pumps make a significant contribution to total carbon export, but there are times when they work in tandem and other times when they work in opposition. The vertical structure of particulate organic carbon is identified as a key target for future observations to better constrain the biological pump. Key Points: Southern Ocean glider and float observations are used to estimate variations in carbon export due to mixed layer and eddy subduction pumpsExport regimes occur when the mixed layer and eddy subduction pumps are in opposition and others when they function in tandemMesoscale coherent structures shape the distribution of submesoscale contributions to the physical carbon pumps [ABSTRACT FROM AUTHOR]

Microbial community dynamics on sinking particles control the amount of carbon that reaches the deep ocean and the length of time that carbon is stored, with potentially profound impacts on Earths climate. A mechanistic understanding of the controls on sinking particle distributions has been hindered by limited depth- and time-resolved sampling and methods that cannot distinguish individual particles. Here, we analyze microbial communities on nearly 400 individual sinking particles in conjunction with more conventional composite particle samples to determine how particle colonization and community assembly might control carbon sequestration in the deep ocean. We observed community succession with corresponding changes in microbial metabolic potential on the larger sinking particles transporting a significant fraction of carbon to the deep sea. Microbial community richness decreased as particles aged and sank; however, richness increased with particle size and the attenuation of carbon export. This suggests that the theory of island biogeography applies to sinking marine particles. Changes in POC flux attenuation with time and microbial community composition with depth were reproduced in a mechanistic ecosystem model that reflected a range of POC labilities and microbial growth rates. Our results highlight microbial community dynamics and processes on individual sinking particles, the isolation of which is necessary to improve mechanistic models of ocean carbon uptake.

In order to quantify pelagic‐benthic coupling on high‐latitude shelves, it is imperative to identify the different physical mechanisms by which phytoplankton are exported to the sediments. In June–July 2023, a field program documented the evolution of an under‐ice phytoplankton bloom on the northeast Chukchi shelf. Here, we use in situ data from the cruise, a simple numerical model, historical water column data, and ocean reanalysis fields to characterize the physical setting and describe the dynamically driven vertical export of chlorophyll associated with the bloom. A water mass front separating cold, high‐nutrient winter water in the north and warmer summer waters to the south—roughly coincident with the ice edge—supported a baroclinic jet which is part of the Central Channel flow branch that veers eastward toward Barrow Canyon. A plume of high chlorophyll fluorescence extending from the near‐surface bloom in the winter water downwards along the front was measured throughout the cruise. Using a passive tracer to represent phytoplankton in the model, it was demonstrated that the plume is the result of subduction due to baroclinic instability of the frontal jet. This process, in concert with the gravitational sinking, pumps the chlorophyll downwards an order of magnitude faster than gravitational sinking alone. Particle tracking using the ocean reanalysis fields reveals that a substantial portion of the chlorophyll away from the front is advected off of the northeast Chukchi shelf before reaching the bottom. This highlights the importance of the frontal subduction process for delivering carbon to the sea floor. Plain Language Summary: The Chukchi Sea shelf north of Bering Strait is known to experience some of the largest phytoplankton blooms in the Arctic Ocean. In 2023, a field program was carried out to quantify aspects of the early summer bloom, with an emphasis on characterizing how the phytoplankton biomass from the bloom is exported to the sea floor. A large bloom was measured under the pack ice in very cold, high‐nutrient water, just north of warmer, ice‐free waters. The front separating the warm and cold waters supported a current flowing eastward, which is one of the main flow pathways on the Chukchi shelf. A plume of high chlorophyll fluorescence extending from the near‐surface bloom downwards along the front was measured throughout the cruise. We demonstrate that this vertical pumping was due to a dynamical process associated with the current which resulted in much faster downward export of phytoplankton than gravitational sinking alone. Tracking the fate of particles on the northeast Chukchi shelf using an ocean simulation revealed that much of the phytoplankton biomass away from the front is carried off the shelf before reaching the bottom. This highlights the importance of the frontal process for delivering chlorophyll to the sea floor. Key Points: An under‐ice phytoplankton bloom developed during June–July in the northeast Chukchi Sea within the Central Channel flow branchA plume of chlorophyll fluorescence extending downwards from the bloom along the current's water mass front was continually presentA simple numerical model demonstrates that the plume is the result of baroclinic instability of the frontal jet [ABSTRACT FROM AUTHOR]

Carbon export driven by submesoscale, eddy‐associated vertical velocities ("eddy subduction"), and particularly its seasonality, remains understudied, leaving a gap in our understanding of ocean carbon sequestration. Here, we assess mechanisms controlling eddy subduction's spatial and seasonal patterns using 15 years of observations from BGC‐Argo floats in the Southern Ocean. We identify signatures of eddy subduction as subsurface anomalies in temperature‐salinity and oxygen. The anomalies are spatially concentrated near weakly stratified areas and regions with strong lateral buoyancy gradients diagnosed from satellite altimetry, particularly in the Antarctic Circumpolar Current's standing meanders. We use bio‐optical ratios, specifically the chlorophyll a to particulate backscatter ratio (Chl/bbp) to find that eddy subduction is most active in the spring and early summer, with freshly exported material associated with seasonally weak vertical stratification and increasing surface biomass. Climate change is increasing ocean stratification globally, which may weaken eddy subduction's carbon export potential. Plain Language Summary: Oceans play an important role in global climate by soaking up and sequestering atmospheric carbon dioxide. Photosynthetic activity at the surface turns carbon dioxide into organic carbon, and if this carbon leaves the surface to the deep ocean, it can be locked away from the atmosphere. One way this occurs is through the physical circulation associated with swirling eddies, which can rapidly transport organic carbon‐rich surface waters and "inject" them into deep waters. However, we still don't fully understand the seasonal timing of this process, or what drives its spatial distribution. We investigated this in the Southern Ocean, which is very important to global climate, using data collected by drifting robots. We find that this process is the most active in regions where eddies drive strong surface stirring, and during the spring, when weak stratification allows injections to penetrate deep into the ocean. Because this process is poorly represented in climate models, these findings will improve our understanding of how the ocean absorbs carbon. Key Points: Eddy subduction in the Southern Ocean is observed as subsurface anomalies in spice and oxygen measured by autonomous profiling floatsSpatial distribution is controlled by weak stratification and strong lateral buoyancy gradients, diagnosed using satellite altimetryBio‐optical proxies suggest that eddy subduction is most active in spring/early summer, driven by weak vertical stratification [ABSTRACT FROM AUTHOR]

Dissolved strontium (Sr) concentrations from Southern Ocean water samples and Sr export fluxes from sediment trap moorings at 1,000 m were used to assess particulate organic carbon (POC) export associated with Acantharia for 2010, 2018, and 2020. The dissolved Sr data revealed a prominent vertical gradient with lower surface Sr concentrations depleted up to 1.5% relative to deep waters. A strong latitudinal surface gradient was observed, ranging from 86.8 μmol kg−1 near the northern end to 88.0 μmol kg−1 near the southern end of a transect through the Australian sector of the Southern Ocean. The vertical and latitudinal gradients are associated with celestite (SrSO4) precipitation by Acantharia and subsequent export to depth. Seasonal variability in Sr export fluxes can be large, particularly during intense events in summer, and reaches a maximum of 11.7 mmol Sr m−2 yr−1, contributing up to 7% of the POC export flux. The coincidence of Sr flux with the second peak of POC export flux implies an association of Acantharia biomass with summertime productivity. Plain Language Summary: Acantharia are tiny marine organisms found worldwide. Their skeletons, composed of a heavy mineral called strontium sulfate (SrSO4, or celestite), make Acantharia important in the oceanic strontium cycling. They also form cysts by resorbing their skeletons and main cellular components, allowing them to sink quickly to depths for reproduction. Despite their dense skeletons and cysts, which could act as effective ballasts, Acantharia were previously thought to have little impact on carbon export in the deeper ocean layers due to their high solubility in seawater. Our study focused on understanding how Acantharia influence strontium cycling and contribute to carbon export in the Southern Ocean. We discovered that Acantharia efficiently remove strontium from the ocean's surface and release it at intermediate depths. These organisms exhibit seasonal variations and are particularly abundant in summer, contributing up to 7% to POC export during the summer productive season. This is significant for the Southern Ocean as a major region for carbon sequestration and global climate buffering. Key Points: Strong vertical and latitudinal gradients of dissolved Sr observed in the Southern Ocean were driven by AcanthariaSr export fluxes exhibited seasonal variations with peaks occurring annually in summer, contributing up to 7% to particulate organic carbon fluxElevated Acantharia biomass is possibly associated with summertime productivity [ABSTRACT FROM AUTHOR]

Marine mesozooplankton play an important role for marine ecosystem functioning and global biogeochemical cycles. Their size structure, varying spatially and temporally, heavily impacts biogeochemical processes and ecosystem services. Mesozooplankton exhibit size changes throughout their life cycle, affecting metabolic rates and functional traits. Despite this variability, many models oversimplify mesozooplankton as a single, unchanging size class, potentially biasing carbon flux estimates. Here, we include mesozooplankton ontogenetic growth and reproduction into a 3‐dimensional global ocean biogeochemical model, PISCES‐MOG, and investigate the subsequent effects on simulated mesozooplankton phenology, plankton distribution, and organic carbon export. Utilizing an ensemble of statistical predictive models calibrated with a global set of observations, we generated monthly climatologies of mesozooplankton biomass to evaluate the simulations of PISCES‐MOG. Our analyses reveal that the model and observation‐based biomass distributions are consistent (rpearson ${\mathrm{r}}_{\mathit{pearson}}$ = 0.40, total epipelagic biomass: 137 TgC from observations vs. 232 TgC in the model), with similar seasonality (later bloom as latitude increases poleward). Including ontogenetic growth in the model induced cohort dynamics and variable seasonal dynamics across mesozooplankton size classes and altered the relative contribution of carbon cycling pathways. Younger and smaller mesozooplankton transitioned to microzooplankton in PISCES‐MOG, resulting in a change in particle size distribution, characterized by a decrease in large particulate organic carbon (POC) and an increase in small POC generation. Consequently, carbon export from the surface was reduced by 10%. This study underscores the importance of accounting for ontogenetic growth and reproduction in models, highlighting the interconnectedness between mesozooplankton size, phenology, and their effects on marine carbon cycling. Key Points: Incorporating mesozooplankton growth and reproduction alters carbon cycling pathways, reducing carbon export at 100 m by 10%Cohort dynamics lead to significant variations in seasonal dynamics across mesozooplankton size classes without affecting export seasonalityStatistical predictive models demonstrate consistency between modeled and observed mesozooplankton dynamics globally [ABSTRACT FROM AUTHOR]

Through biological activity, marine dissolved inorganic carbon (DIC) is transformed into different types of biogenic carbon available for export to the ocean interior, including particulate organic carbon (POC), dissolved organic carbon (DOC), and particulate inorganic carbon (PIC). Each biogenic carbon pool has a different export efficiency that impacts the vertical ocean carbon gradient and drives natural air-sea carbon dioxide gas (CO2) exchange. In the Southern Ocean (SO), which presently accounts for ~40% of the anthropogenic ocean carbon sink, it is unclear how the production of each biogenic carbon pool contributes to the contemporary air-sea CO2 exchange. Based on 107 independent observations of the seasonal cycle from 63 biogeochemical profiling floats, we provide the basin-scale estimate of distinct biogenic carbon pool production. We find significant meridional variability with enhanced POC production in the subantarctic and polar Antarctic sectors and enhanced DOC production in the subtropical and sea-ice-dominated sectors. PIC production peaks between 47°S and 57°S near the great calcite belt. Relative to an abiotic SO, organic carbon production enhances CO2 uptake by 2.80 ± 0.28 Pg C y-1, while PIC production diminishes CO2 uptake by 0.27 ± 0.21 Pg C y-1. Without organic carbon production, the SO would be a CO2 source to the atmosphere. Our findings emphasize the importance of DOC and PIC production, in addition to the well-recognized role of POC production, in shaping the influence of carbon export on air-sea CO2 exchange.

Eroded soils sustain a substantial part of organic matter in tidal rivers adjacent to estuaries, and photochemical transformations of soils in tidal rivers would influence estuarine elemental cycles. However, complex aquatic environments and diverse soil sources complicate the enrichment of dissolved organic matter (DOM) photoreleased from soils. Here, we conducted a 7-day irradiation experiment for seven kinds of soils from the lower basin of Dagu River (DGR) in the laboratory to study the influence of salinity and soil properties on DOM chemistry by characterizing the content and optical properties of DOM. Results showed that light cultures had higher amount of DOM and humic-like components than dark cultures. Principal component analysis (PCA) and Mantel's analysis found that salinity and soil properties significantly influence the production of photoreleased DOM, especially humic-like components. Salinity could inhibit the photodissolution of soils, and aged soils with low δ13CSOM released more DOM and humic-like components. Although the DGR is impacted by intruded seawater, high content of photoreleased DOM in seawater cultures still pointed out the important contribution of soil photodissolution to the DOM reservoir of tidal rivers. Considering high proportion of humic-like components in photoreleased DOM, photochemical transformations of soils in tidal rivers would promote the export flux of carbon from estuaries to open seas. This study emphasizes the importance of soil photodissolution of tidal rivers in the carbon transfer from lands to oceans. [ABSTRACT FROM AUTHOR]

Vertical migrants are a diverse group of organisms, which includes crustaceans, cephalopods and mesopelagic fishes. They play an active role in the biogeochemical cycles but are in general not included in numerical models. In this study we introduce a fully coupled Earth system model that represents vertical migration and with this resolves the key components of the mesopelagic ecosystem, namely migrating zooplankton and mesopelagic fish, including their feedbacks on biogeochemical cycles. The redistribution of nutrients in the water column by vertical migration results in a reduction of the net primary production of 14%–21%, as well as in an asymmetric response in the low oxygenated waters in the tropical Pacific (an increase in the northern and a decrease in the southern oxygen minimum zone). On a global scale, we find the active transport of carbon out of the surface layer to be equivalent to ∼25% of the total export (∼30% relative to passive sinking). In the low latitudes, migration results regionally in a reduction of the shallow export by 2%–10% and an increase of the deep carbon export by 6%–15%. In our simulations, mesopelagic fish, with a biomass of 3–3.4 Gt wet weight, have a slightly larger impact on active carbon flux than migrating zooplankton. Key Points: We present a global model that represents the impact of mesopelagic fish and vertically migrating organisms on the biogeochemical cyclesThe simulated redistribution of nutrients by vertical migration results in a reduction of the global net primary production by 14%–21% compared to the simulation without vertical migrationThe active carbon export out of the surface layers by vertical migration amounts to ∼25% of the total transport (∼30% of the passive sinking) [ABSTRACT FROM AUTHOR]

The North Atlantic is a region of enhanced biogeochemical and climatological importance for the global ocean as it is the site of one of the largest seasonal phytoplankton blooms on the planet. However, there is a lack of understanding of how phytoplankton size influences bloom dynamics and associated nutrient utilization rates, particularly during the decline phase when export to the deep ocean is especially pronounced. Here, we evaluate trends in size‐fractionated carbon, nitrogen, and silicic acid uptake rates in conjunction with environmental parameters to assess these dynamics. In our study, the decline phase of the bloom continued to be highly productive with net primary production (NPP) ranging from 36.4 to 146.6 mmol C m−2 d−1 and approximately 54% of primary production being driven by large phytoplankton cells (≥5 μm) that were primarily utilizing nitrate (mean f‐ratio of 0.77). Entrainment of silicic acid related to deepening of the mixed layer caused by storms increased silicic acid uptake rates to 2.0–5.7 mmol Si m−2 d−1 without concomitant increases in NPP by large cells (silicic acid to carbon uptake ratios averaged 0.12). A companion study in the North Pacific allowed for paired evaluation of these regions. Our results suggest that in highly productive regions where phytoplankton biomass and productivity is distributed across a broad range of cell sizes, such as the North Atlantic, size itself has a stronger influence on nutrient cycling and potential carbon export relative to regions with lower production and a predominance of small (<5 μm) cells, such as the North Pacific. Plain Language Summary: The North Atlantic Ocean experiences a seasonal bloom of phytoplankton. This bloom represents one of the largest removals of anthropogenic carbon to the deep ocean on the planet. Here, we seek to better quantify the mechanisms of this removal by characterizing the amount and composition of phytoplankton present and how much primary production they are engaging in. We sampled during the decline phase of the spring bloom and found the region to be characterized by mostly large phytoplankton primarily utilizing nitrate. When compared to a companion study in the North Pacific, this study confirms the important role cell size and nutrient availability play in determining the degree of productivity in both high and low primary production oceanic regions. Key Points: The decline phase of the North Atlantic spring bloom exhibits transitions in the balance between small and large size‐fractionated primary production dynamicsDuring the bloom decline, diatoms contribute less to primary production but remain a substantial component of phytoplankton biomassPhytoplankton size is a substantial control on ecosystem dynamics in endmember oceanic productivity‐export systems [ABSTRACT FROM AUTHOR]

This study analyzes the molecular composition of dissolved organic matter (DOM) in Prydz Bay by Fourier Transform Ion Cyclotron Resonance mass spectrometry to probe the carbon sequestration capacity in the continental shelf system. Concentrations of particulate organic carbon (POC), particulate nitrogen and dissolved organic carbon (DOC) with water depth show that POC could be mainly decomposed into DOC and/or microbially degraded. Highly labile DOC is further degraded and remineralized by microorganisms within the upper 200 m, as evidenced by a downward enrichment of 13CPOC and increases in the average molecular weight, oxygen atom number (O) and double bond equivalents of DOM molecules, indicating that biodegradation is the main driver for particulate organic matter and DOM evolution with water depth. Semi‐quantitative calculation demonstrates that ∼83% of POC was transformed to DOC as well as dissolved inorganic carbon (DIC), and ∼30% of DOC further to DIC via microbial degradation within the upper 200 m in summer, resulting in a relatively low total organic carbon content in sediments of Prydz Bay. The newly transformed DIC and residue DOC can be preserved in the deep layer due to the formation of well stratified and stable water body in summer of Prydz Bay, ultimately entering the regional circulation system instead of being released back into the atmosphere. This could be one of the most important processes determining the atmosphere CO2 uptake in the continental shelf system of Southern Ocean. Plain Language Summary: The continental shelf system in the Southern Ocean, especially with the development of polynyas, plays a critical role in the global carbon cycle and disproportionally accounts for almost half of the anthropogenic CO2 uptake by the biological pump, with ensuing deep‐sea sequestration by the form of particulate and dissolved organic carbon, and even CO2 due to microbial degradation of primary productivity. However, complex interactions between physical, chemical and biological processes significantly affect the fate of organic carbon through the water column, resulting in an uncertainty of vertical carbon export. In this study, we conduct an investigation on the dissolved organic matter sources and evolution with water depth in summer of Prydz Bay, a typical polynya‐developed continental shelf system in the East Antarctica. The results clearly show that biodegradation is the main factor controlling organic carbon mineralization and mainly occurs within the upper 200 m of water column, with a transformation into dissolved organic carbon (DOC) and dissolved inorganic carbon (DIC). Integrated with regional circulation system and hydrological characteristics, our study further indicates that newly formed DIC and DOC can be preserved in the deep layer of Prydz Bay, then into the regional circulation system rather back to the atmosphere. Key Points: Novel evidence at molecular level shows the vital role of biodegradation during carbon sink in Prydz BayThe conversion of POC into DOC/DIC mainly occurs within the upper 200 m of water columnHighlighting regional circulation's contribution to carbon transport and sequestration in the Southern Ocean [ABSTRACT FROM AUTHOR]

We use observations from the Southern Ocean (SO) biogeochemical profiling float array to quantify the meridional pattern of particle export efficiency (PEeff) during the austral productive season. Float estimates reveal a pronounced latitudinal gradient of PEeff, which is quantitatively supported by a compilation of existing ship‐based measurements. Relying on complementary float‐based estimates of distinct carbon pools produced through biological activity, we find that PEeff peaks near the region of maximum particulate inorganic carbon sinking flux in the polar antarctic zone, where net primary production (NPP) is the lowest. Regions characterized by intermediate NPP and low PEeff, primarily in the subtropical and seasonal ice zones, are generally associated with a higher fraction of dissolved organic carbon production. Our study reveals the critical role of distinct biogenic carbon pool production in driving the latitudinal pattern of PEeff in the SO. Plain Language Summary: Microbial organisms in seawater transform carbon dioxide into different types of carbon through photosynthesis and food web cycling. These carbon types include particulate and dissolved phases, with particles being more efficiently transferred out of the sunlit ocean via gravitational sinking. The ratio of sinking particulate organic carbon to total organic carbon production, commonly referred to as the particle export efficiency, is a metric used to describe how efficiently carbon moves from the surface to the deep ocean. Using observations from a large array of robots in the Southern Ocean, we find that the different types of biogenic carbon produced control the latitudinal gradient in particle export efficiency, which is highest in regions where particulate inorganic carbon export is greatest, even when photosynthetically fixed carbon is minimal. In other areas where phytoplankton carbon production is moderate but largely comprised of dissolved organic carbon, the particle export efficiency is lower. Key Points: Meridional pattern of particle export efficiency (PEeff) estimated from BGC‐Argo aligns with ship‐based observations in the Southern OceanLow PEeff in subtropical and ice‐covered regions and high PEeff in subpolar regions is linked to the biogenic carbon pools producedMost global models struggle to reproduce the meridional pattern of PEeff in the Southern Ocean [ABSTRACT FROM AUTHOR]

The demand for marine living resources is increasing at an unprecedented scale because of the need for continuous food provision to the world’s population. The potential of already exploited fish stocks to meet this demand is limited. Therefore, mesopelagic fish have recently become attractive potential targets for fisheries because of their vast conjectured biomass. However, the role of mesopelagic fish in marine ecosystems is poorly understood. Before developing commercial exploitation plans, the relationship between mesopelagic fish and other groups in the marine food web and biogeochemical cycles should be analyzed quantitatively. In this study, we coupled a one-dimensional biogeochemical model (North Atlantic Generic Ecosystem Model) with a higher-trophic-level food web model (Ecopath with Ecosim) for the Sargasso Sea in the North Atlantic to investigate changes in carbon export and trophodynamics under two mesopelagic fish harvesting scenarios. The coupled model represented the marine food web from plankton to fish and mammals, vertical carbon export dynamics, and their interaction with fisheries. The results showed that when mesopelagic fish were not harvested, they contributed approximately 6% of the total carbon export in the surface waters, but up to 40% of the total carbon export below 400 m. Harvesting mesopelagic fish altered the energy transfers within the food web as well as to fisheries. The ecological footprint of fisheries increased significantly. Due to declining competition in the food web, epipelagic fish increased to exert elevated grazing pressure on phytoplankton; hence, phytoplankton-mediated carbon export decreased. The total carbon export decreased by 14% due to the decreases in mesopelagic fish- and phytoplanktonmediated carbon exports. The simulated increase in zooplankton- and nonmesopelagic fish-mediated carbon exports (up to 92% and 96%, respectively) did not compensate for the total decrease in carbon exports under harvesting scenarios. The findings of this study highlighted that mesopelagic fish not only have a direct control on carbon dynamics by their metabolic releases and diel vertical migration, but also strong indirect controls through prey-predator interactions within the food web. Therefore, the implications of harvesting mesopelagic fish should be carefully considered from a holistic perspective. [ABSTRACT FROM AUTHOR]

Dissolved organic and inorganic carbon (DOC and DIC) influence water quality, ecosystem health, and carbon cycling. Dissolved carbon species are produced by biogeochemical reactions and laterally exported to streams via distinct shallow and deep subsurface flow paths. These processes are arduous to measure and challenge the quantification of global carbon cycles. Here we ask: when, where, and how much is dissolved carbon produced in and laterally exported from the subsurface to streams? We used a catchment‐scale reactive transport model, BioRT‐HBV, with hydrometeorology and stream carbon data to illuminate the "invisible" subsurface processes at Sleepers River, a carbonate‐based catchment in Vermont, United States. Results depict a conceptual model where DOC is produced mostly in shallow soils (3.7 ± 0.6 g/m2/yr) and in summer at peak root and microbial respiration. DOC is flushed from soils to the stream (1.0 ± 0.2 g/m2/yr) especially during snowmelt and storms. A large fraction of DOC (2.5 ± 0.2 g/m2/yr) percolates to the deeper subsurface, fueling deep respiration to generate DIC. DIC is exported predominantly from the deeper subsurface (7.1 ± 0.4 g/m2/yr, compared to 1.3 ± 0.3 g/m2/yr from shallow soils). Deep respiration reduces DOC and increases DIC concentrations at depth, leading to commonly observed DOC flushing (increasing concentrations with discharge) and DIC dilution patterns (decreasing concentrations with discharge). Surprisingly, respiration processes generate more DIC than weathering in this carbonate‐based catchment. These findings underscore the importance of vertical connectivity between the shallow and deep subsurface, highlighting the overlooked role of deep carbon processing and export. Plain Language Summary: Dissolved organic and inorganic carbon (DOC and DIC) are important chemical species that affect water quality, ecosystem health, and carbon dioxide emissions from streams. DOC and DIC are produced through different reactions at and below the ground surface before they are transported to streams through underground flow paths. However, it is difficult to measure and observe these reactions and transport pathways, limiting our understanding of when, where, and how much dissolved carbon species are produced and exported from distinct subsurface depths. Here we used a computational model, BioRT‐HBV, to simulate reactions and transport processes and to better understand the production and export of dissolved carbon at Sleepers River, a small catchment in Vermont, United States. Results show that DOC was primarily produced through shallow subsurface reactions and exported through shallow flow paths. DIC was produced in both the shallow and deep subsurface but primarily exported through deep flow paths. Reactions that produced DOC and DIC occurred faster under warm and wet conditions (summer and spring), while export of DOC and DIC increased under wet conditions (spring, snowmelt, storms). Results suggest that climate change may affect the production and export of dissolved carbon species through increased temperatures and intense storm events. Key Points: Dissolved organic carbon (DOC) was mainly produced in warm summer and exported in wet spring from shallow subsurfaceDissolved inorganic carbon (DIC) was comparably produced in the shallow and deep subsurface but exported mostly from the deep subsurface in wet springDIC originated more from biogenic (soil respiration and deep respiration) than geogenic (carbonate weathering) sources in a carbonate‐based catchment [ABSTRACT FROM AUTHOR]

A large proportion of algal production and carbon export occurs after snowmelt and before ice melt in the Arctic Ocean. To determine the magnitude of under-ice export fluxes over the Chukchi Plateau (CP), a total of 25 sediment trap deployments were completed at two ice camps during the annual field survey of the Korean IBRV Araon in 2018: a first ice camp (CP1) conducted from August 17 to 19 and a second ice camp (CP2) conducted from August 20 to 22. Chlorophyll a (chl a) and particulate organic carbon (POC) fluxes were measured at 2, 5, 10 and 30 m under ice, and zooplankton collected in the sediment traps were enumerated and identified. Both chl a and POC fluxes were the highest at 5 m under ice at CP2, likely due to enhanced fluxes of the sea ice algae Melosira arctica and to a potential higher release of particulate matter from a thinner sea ice cover. Whereas Calanus glacialis/marshallae were dominant at all depths and both sites, the large numbers of individuals observed at 10 m at CP1 likely enhanced zooplankton grazing pressure at that site, further contributing to the high spatial variability in under-ice export fluxes. Overall, under-ice fluxes obtained in August 2018 highlighted the importance of M. arctica aggregates as a source of carbon for pelagic consumers and for carbon export during summer over the Chukchi Plateau. [ABSTRACT FROM AUTHOR]

The demand for marine living resources is increasing at an unprecedented scale because of the need for continuous food provision to the world’s population. The potential of already exploited fish stocks to meet this demand is limited. Therefore, mesopelagic fish have recently become attractive potential targets for fisheries because of their vast conjectured biomass. However, the role of mesopelagic fish in marine ecosystems is poorly understood. Before developing commercial exploitation plans, the relationship between mesopelagic fish and other groups in the marine food web and biogeochemical cycles should be analyzed quantitatively. In this study, we coupled a one-dimensional biogeochemical model (North Atlantic Generic Ecosystem Model) with a higher-trophic-level food web model (Ecopath with Ecosim) for the Sargasso Sea in the North Atlantic to investigate changes in carbon export and trophodynamics under two mesopelagic fish harvesting scenarios. The coupled model represented the marine food web from plankton to fish and mammals, vertical carbon export dynamics, and their interaction with fisheries. The results showed that when mesopelagic fish were not harvested, they contributed approximately 6% of the total carbon export in the surface waters, but up to 40% of the total carbon export below 400 m. Harvesting mesopelagic fish altered the energy transfers within the food web as well as to fisheries. The ecological footprint of fisheries increased significantly. Due to declining competition in the food web, epipelagic fish increased to exert elevated grazing pressure on phytoplankton; hence, phytoplankton-mediated carbon export decreased. The total carbon export decreased by 14% due to the decreases in mesopelagic fish- and phytoplankton-mediated carbon exports. The simulated increase in zooplankton- and non-mesopelagic fish-mediated carbon exports (up to 92% and 96%, respectively) did not compensate for the total decrease in carbon exports under harvesting scenarios. The findings of this study highlighted that mesopelagic fish not only have a direct control on carbon dynamics by their metabolic releases and diel vertical migration, but also strong indirect controls through prey-predator interactions within the food web. Therefore, the implications of harvesting mesopelagic fish should be carefully considered from a holistic perspective.

The 13C isotope composition (δ13C) of leaf dry matter is a useful tool for physiological and ecological studies. However, how post-photosynthetic fractionation associated with respiration and carbon export influences δ13C remains uncertain. We investigated the effects of post-photosynthetic fractionation on δ13C of mature leaves of Cleistogenes squarrosa , a perennial C4 grass, in controlled experiments with different levels of vapour pressure deficit and nitrogen supply. With increasing leaf age class, the 12C/13C fractionation of leaf organic matter relative to the δ13C of atmosphere CO2 (ΔDM) increased while that of cellulose (Δcel) was almost constant. The divergence between ΔDM and Δcel increased with leaf age class, with a maximum value of 1.6‰, indicating the accumulation of post-photosynthetic fractionation. Applying a new mass balance model that accounts for respiration and export of photosynthates, we found an apparent 12C/13C fractionation associated with carbon export of –0.5‰ to –1.0‰. Different ΔDM among leaves, pseudostems, daughter tillers, and roots indicate that post-photosynthetic fractionation happens at the whole-plant level. Compared with ΔDM of old leaves, ΔDM of young leaves and Δcel are more reliable proxies for predicting physiological parameters due to the lower sensitivity to post-photosynthetic fractionation and the similar sensitivity in responses to environmental changes. [ABSTRACT FROM AUTHOR]

Diazotrophs regulate marine productivity in the oligotrophic ocean by alleviating nitrogen limitation, contributing to particulate organic carbon (POC) export to the deep ocean. Yet, the characterization of particles composing the sinking POC flux has never been explored in such ecosystems. Moreover, the contribution of the direct gravitational export of diazotrophs to the overall flux is seldom assessed. Here we explore the composition of the sinking POC flux in a hot spot of N2 fixation (the western sub-tropical South Pacific) using polyacrylamide gel-filled traps deployed at two stations (S05M and S10M) and three depths (170 m, 270 m, 1000 m) during the TONGA expedition (November-December 2019). Image analyses of particles collected in the gels was used to classify them into 5 categories (fecal aggregates, phytodetrital aggregates, mixed aggregates, cylindrical fecal pellets, and zooplankton carcasses). Fecal aggregates were the most abundant at both stations and all depths and dominated the flux (average of 56 ± 28% of the POC flux), followed by zooplankton carcasses (24 ± 19%), cylindrical fecal pellets (15 ± 14%) and mixed aggregates (5 ± 4%), whereas phytodetrital aggregates contributed less (<1%). Since N isotope budgets show that export is mainly supported by diazotrophy at these stations, these results suggest that the diazotroph-derived N has been efficiently transferred to the foodweb up to zooplankton and fecal pellets before being exported, pleading for an indirect export of diazotrophy. However, random confocal microscopy examination performed on sinking particles revealed that diazotrophs were present in several categories of exported particles, suggesting that diazotrophs are also directly exported, with a potential contribution to overall POC fluxes increasing with depth. Our results provide the first characterization of particle categories composing the sinking flux and their contribution to the overall flux in a hot spot of N2 fixation. [ABSTRACT FROM AUTHOR]

Mesozooplankton is a very diverse group of small animals ranging in size from 0.2 to 20 mm not able to swim against ocean currents. It is a key component of pelagic ecosystems through its roles in the trophic networks and the biological carbon pump. Traditionally studied through microscopes, recent methods have been however developed to rapidly acquire large amounts of data (morphological, molecular) at the individual scale, making it possible to study mesozooplankton using a trait‐based approach. Here, combining quantitative imaging with metabarcoding time‐series data obtained in the Sargasso Sea at the Bermuda Atlantic Time‐series Study (BATS) site, we showed that organisms' transparency might be an important trait to also consider regarding mesozooplankton impact on carbon export, contrary to the common assumption that just size is the master trait directing most mesozooplankton‐linked processes. Three distinct communities were defined based on taxonomic composition, and succeeded one another throughout the study period, with changing levels of transparency among the community. A co‐occurrences' network was built from metabarcoding data revealing six groups of taxa. These were related to changes in the functioning of the ecosystem and/or in the community's morphology. The importance of Diel Vertical Migration at BATS was confirmed by the existence of a group made of taxa known to be strong migrators. Finally, we assessed if metabarcoding can provide a quantitative approach to biomass and/or abundance of certain taxa. Knowing more about mesozooplankton diversity and its impact on ecosystem functioning would allow to better represent them in biogeochemical models. [ABSTRACT FROM AUTHOR]

Introduction: Artificial upwelling has been discussed as a nature-based solution to fertilize currently unproductive areas of the ocean to enhance food web productivity and atmospheric CO2 sequestration. The efficacy of this approach may be closely tied to the nutrient stoichiometry of the upwelled water, as Si- rich upwelling should benefit the growth of diatoms, who are key players for primary production, carbon export and food web efficiency. Methods: With a mesocosm experiment in subtropical waters, we assessed the physiological and functional responses of an oligotrophic phytoplankton community to artificial upwelling under varying Si:N ratios (0.07-1.33). Results: Deep water fertilization led to strongly enhanced primary productivity rates and net autotrophy across Si scenarios. At the community level, Si-rich upwelling50 temporarily increased primary production and consistently enhanced diatom growth, producing up to 10-fold higher abundances compared to Si-deficient upwelling. At the organism level, contrasting effects were observed. On the one hand, silicification and size of diatom cells remained unaffected by Si:N, which is surprising given the direct dependency of these traits on Si. On the other hand, diatom Chlorophyll a density and carbon density were strongly reduced and particulate matter C:N was elevated under Si- rich upwelling. Discussion: This suggests a reduced nutritional value for higher trophic levels under high Si:N ratios. Despite these strong qualitative changes under high Si, diatom cells appeared healthy and showed high photosynthetic efficiency. Our findings reveal great physiological plasticity and adaptability in phytoplankton under artificial upwelling, with Si-dependent trade-offs between primary producer quantity and quality. [ABSTRACT FROM AUTHOR]

IntroductionArtificial upwelling has been discussed as a nature-based solution to fertilize currently unproductive areas of the ocean to enhance food web productivity and atmospheric CO2 sequestration. The efficacy of this approach may be closely tied to the nutrient stoichiometry of the upwelled water, as Si-rich upwelling should benefit the growth of diatoms, who are key players for primary production, carbon export and food web efficiency.MethodsWith a mesocosm experiment in subtropical waters, we assessed the physiological and functional responses of an oligotrophic phytoplankton community to artificial upwelling under varying Si:N ratios (0.07-1.33).ResultsDeep water fertilization led to strongly enhanced primary productivity rates and net autotrophy across Si scenarios. At the community level, Si-rich upwelling50 temporarily increased primary production and consistently enhanced diatom growth, producing up to 10-fold higher abundances compared to Si-deficient upwelling. At the organism level, contrasting effects were observed. On the one hand, silicification and size of diatom cells remained unaffected by Si:N, which is surprising given the direct dependency of these traits on Si. On the other hand, diatom Chlorophyll a density and carbon density were strongly reduced and particulate matter C:N was elevated under Si-rich upwelling.DiscussionThis suggests a reduced nutritional value for higher trophic levels under high Si:N ratios. Despite these strong qualitative changes under high Si, diatom cells appeared healthy and showed high photosynthetic efficiency. Our findings reveal great physiological plasticity and adaptability in phytoplankton under artificial upwelling, with Si-dependent trade-offs between primary producer quantity and quality.

Diazotrophs regulate marine productivity in the oligotrophic ocean by alleviating nitrogen limitation, contributing to particulate organic carbon (POC) export to the deep ocean. Yet, the characterization of particles composing the sinking POC flux has never been explored in such ecosystems. Moreover, the contribution of the direct gravitational export of diazotrophs to the overall flux is seldom assessed. Here we explore the composition of the sinking POC flux in a hot spot of N2 fixation (the western sub-tropical South Pacific) using polyacrylamide gel-filled traps deployed at two stations (S05M and S10M) and three depths (170 m, 270 m, 1000 m) during the TONGA expedition (November-December 2019). Image analyses of particles collected in the gels was used to classify them into 5 categories (fecal aggregates, phytodetrital aggregates, mixed aggregates, cylindrical fecal pellets, and zooplankton carcasses). Fecal aggregates were the most abundant at both stations and all depths and dominated the flux (average of 56 ± 28% of the POC flux), followed by zooplankton carcasses (24 ± 19%), cylindrical fecal pellets (15 ± 14%) and mixed aggregates (5 ± 4%), whereas phytodetrital aggregates contributed less (

The Antarctic Marginal Ice Zone (MIZ) accounts for 15% of the Southern Ocean's primary production (PP), but limited data has hindered understanding of its variability and connection to carbon export. Using a combination of gliders, biogeochemical Argo floats and satellite observations in the northeast Weddell Sea, we show that years with more sea‐ice formation over winter are followed by more intense phytoplankton blooms (∼15% greater daily PP) and export to 100 m (∼50% higher daily carbon export) the following summer. However, the carbon export beyond the deepest winter mixed layer did not vary in proportion to PP, suggesting different drivers of carbon export at depth compared to surface waters. Furthermore, across the entire MIZ, the response of blooms to sea‐ice volume was spatially variable, indicating the need to consider spatial heterogeneity in the response of the biological carbon pump to future sea‐ice changes. Plain Language Summary: Algae in the ocean surface take up carbon dioxide from the atmosphere through photosynthesis and transfer it to the deep ocean when they die and sink. This process is key to maintaining a habitable planet and is known as the biological carbon pump (BCP). The seasonally ice‐covered ocean around Antarctica is one of the most active areas for algal growth, but also a region of rapid climate change. Because of the difficulty in taking measurements in this remote region, the physical and biological processes that control the growth and sinking of algae and its response to changing sea‐ice remain uncertain. In this study, we use a combination of satellites and autonomous robots to elucidate the role of sea‐ice variability on the BCP. We find that sea‐ice impacts algal growth by its influence on both the light and nutrient conditions needed for photosynthesis. Predicting the amount of algae that subsequently sinks to depth as carbon flux, although influenced by sea‐ice conditions, is more complex and linked to the greater marine ecosystem. Evidence suggests that the species of algae, zooplankton grazing, and the rate at which dead algae breaks down and sinks are important and should be a focus point for further research. Key Points: High‐resolution in‐situ observations are used to characterize multi‐year phytoplankton bloom phenology and amplitude in the Antarctic Marginal Ice ZoneYears with greater sea‐ice volume drive deeper mixing that tend to support higher magnitude blooms in the northeast Weddell SeaCarbon export efficiency is affected by bloom magnitude, community composition and water column stratification [ABSTRACT FROM AUTHOR]

Earth System Models generally predict increasing upper ocean stratification from 21st century warming, which will cause a decrease in the vertical nutrient flux forcing declines in marine net primary productivity (NPP) and carbon export. Recent advances in quantifying marine ecosystem carbon to nutrient stoichiometry have identified large latitudinal and biome variability, with low‐latitude oligotrophic systems harboring pico‐sized phytoplankton exhibiting large phosphorus to carbon cellular plasticity. The climate forced changes in nutrient flux stoichiometry and phytoplankton community composition are thus likely to alter the ocean's biogeochemical response and feedback with the carbon‐climate system. We have added three pico‐phytoplankton functional types within the Biogeochemical Elemental Cycling component of the Community Earth System Model while incorporating variable cellular phosphorus to carbon stoichiometry for all represented phytoplankton types. The model simulates Prochlorococcus and Synechococcus populations that dominate the productivity and sinking carbon export of the tropical and subtropical ocean, and pico‐eukaryote populations that contribute significantly to productivity and export within the subtropical to mid‐latitude transition zone, with the western subtropical regions of each basin supporting the most P‐poor stoichiometries. Subtropical gyre recirculation regions along the poleward flanks of surface western boundary currents are identified as regional hotspots of enhanced carbon export exhibiting C‐rich/P‐poor stoichiometry, preferentially inhabited by pico‐eukaryotes and diatoms. Collectively, pico‐phytoplankton contribute ∼58% of global NPP and ∼46% of global particulate organic carbon export below 100 m through direct and ecosystem processing pathways. Biodiversity and cellular nutrient plasticity in marine pico‐phytoplankton combine to increase their contributions to ocean productivity and the biological carbon pump. Key Points: Simulated Prochlorococcus, Synechococcus, and pico‐eukaryotes contribute ∼60% of marine net primary productivity (NPP)Pico‐phytoplankton cycling contributes half of the marine export production, approaching parity with their contribution to NPPPico‐eukaryotes and diatoms with elevated C:P stoichiometry enhance carbon export at poleward flanks of western boundary currents [ABSTRACT FROM AUTHOR]

How coral reefs with high productivity and biodiversity can flourish in oligotrophic tropical oceans has inspired substantial research on coral reef ecosystems. Increasing evidence shows that similar to water in an oasis in the desert, there are stable nutrient supplies to coral reefs in oligotrophic oceans. Here, with emphasis on the fluxes of organic matter, we summarize at the ecosystem level (1) the multiple input pathways of external nutrients, (2) the storage of nutrients in reef organisms, (3) the efficient retaining and recycling of dissolved and particulate organic matter within coral reef ecosystems, (4) the distinctly high phytoplankton productivity and biomass inside and near oceanic coral reefs, and (5) the export of reef-related organic carbon to adjacent open oceans. These properties enable coral reefs to function as ecological "pumps" for gathering nutrients across ecosystems and space, retaining and recycling nutrients within the ecosystem, supporting high phytoplankton productivity, and exporting organic carbon to adjacent open oceans. Particularly, the high phytoplankton productivity and biomass make waters around coral reefs potential hotspots of carbon export to ocean depths via the biological pump. We demonstrate that organic carbon influx is vital for coral reef ecosystems' carbon budget and carbon export. The concept of the coral reef ecological pump provides a framework to improve the understanding of the functioning of the coral reef ecosystem and its responses to disturbance. Prospects of the coral reef ecological pump in coral reef studies are discussed in changing oceans driven by human activities and global change in the Anthropocene. [ABSTRACT FROM AUTHOR]

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.

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.

Whereas the vertical transport of biomass from productive surface waters to the deep ocean (the biological pump) is a critical component of the global carbon cycle, its magnitude and variability is poorly understood. Global-scale estimates of ocean carbon export vary widely, ranging from ∼5 to ∼20 Gt C y – 1 due to uncertainties in methods and unclear definitions. Satellite-derived properties such as phytoplankton biomass, sea surface temperature, and light attenuation at depth provide information about the oceanic ecosystem with unprecedented coverage and resolution in time and space. These products have been the basis of an intense effort over several decades to constrain different biogeochemical production rates and fluxes in the ocean. One critical challenge in this effort has been to estimate the magnitude of the biological pump from satellite-derived properties by establishing how much of the primary production is exported out of the euphotic zone, a flux that is called export production. Here we present a review of existing algorithms for estimating export production from satellite-derived properties, available in-situ datasets that can be used for testing the algorithms, and earlier evaluations of the proposed algorithms. The satellite-derived products used in the algorithm evaluation are all based largely on the Ocean Colour Climate Change Initiative (OC-CCI) products, and carbon products derived from them. The different resources are combined in a meta-analysis.

Whales have been titled climate savers in the media with their recovery welcomed as a potential carbon solution. However, only a few studies were performed to date providing data or model outputs to support the hypothesis. Following an outline of the primary mechanisms by which baleen whales remove carbon from the atmosphere for eventual sequestration at regional and global scales, we conclude that the amount of carbon whales are potentially sequestering might be too little to meaningfully alter the course of climate change. This is in contrast to media perpetuating whales as climate engineers. Creating false hope in the ability of charismatic species to be climate engineers may act to further delay the urgent behavioral change needed to avert catastrophic climate change impacts, which can in turn have indirect consequences for the recovery of whale populations. Nevertheless, whales are important components of marine ecosystems, and any further investigation on existing gaps in their ecology will contribute to clarifying their contribution to the ocean carbon cycle, a major driver of the world’s climate. While whales are vital to the healthy functioning of marine ecosystems, overstating their ability to prevent or counterbalance anthropogenically induced changes in global carbon budget may unintentionally redirect attention from known, well-established methods of reducing greenhouse gases. Large scale protection of marine environments including the habitats of whales will build resilience and assist with natural carbon capture.

ABSTRACT The biological carbon pump (BCP) in the Southern Ocean is driven by phytoplankton productivity and is a significant organic matter sink. However, the role of particle-attached (PA) and free-living (FL) prokaryotes (bacteria and archaea) and their diversity in influencing the efficiency of the BCP is still unclear. To investigate this, we analyzed the metagenomes linked to suspended and sinking marine particles from the Sub-Antarctic Southern Ocean Time Series (SOTS) by deploying a Marine Snow Catcher (MSC), obtaining suspended and sinking particulate material, determining organic carbon and nitrogen flux, and constructing metagenome-assembled genomes (MAGs). The suspended and sinking particle-pools were dominated by bacteria with the potential to degrade organic carbon. Bacterial communities associated with the sinking fraction had more genes related to the degradation of complex organic carbon than those in the suspended fraction. Archaea had the potential to drive nitrogen metabolism via nitrite and ammonia oxidation, altering organic nitrogen concentration. The data revealed several pathways for chemoautotrophy and the secretion of recalcitrant dissolved organic carbon (RDOC) from CO2, with bacteria and archaea potentially sequestering particulate organic matter (POM) via the production of RDOC. These findings provide insights into the diversity and function of prokaryotes in suspended and sinking particles and their role in organic carbon/nitrogen export in the Southern Ocean. IMPORTANCE The biological carbon pump is crucial for the export of particulate organic matter in the ocean. Recent studies on marine microbes have shown the profound influence of bacteria and archaea as regulators of particulate organic matter export. Yet, despite the importance of the Southern Ocean as a carbon sink, we lack comparable insights regarding microbial contributions. This study provides the first insights regarding prokaryotic contributions to particulate organic matter export in the Southern Ocean. We reveal evidence that prokaryotic communities in suspended and sinking particle fractions harbor widespread genomic potential for mediating particulate organic matter export. The results substantially enhance our understanding of the role played by microorganisms in regulating particulate organic matter export in suspended and sinking marine fractions in the Southern Ocean.

Through biological activity, marine dissolved inorganic carbon (DIC) is transformed into different types of biogenic carbon available for export to the ocean interior, including particulate organic carbon (POC), dissolved organic carbon (DOC), and particulate inorganic carbon (PIC). Each biogenic carbon pool has a different export efficiency that impacts the vertical ocean carbon gradient and drives natural air-sea carbon dioxide gas (CO2) exchange. In the Southern Ocean (SO), which presently accounts for ~40% of the anthropogenic ocean carbon sink, it is unclear how the production of each biogenic carbon pool contributes to the contemporary air-sea CO2 exchange. Based on 107 independent observations of the seasonal cycle from 63 biogeochemical profiling floats, we provide the basin-scale estimate of distinct biogenic carbon pool production. We find significant meridional variability with enhanced POC production in the subantarctic and polar Antarctic sectors and enhanced DOC production in the subtropical and sea-ice-dominated sectors. PIC production peaks between 47°S and 57°S near the "great calcite belt." Relative to an abiotic SO, organic carbon production enhances CO2 uptake by 2.80 ± 0.28 Pg C y-1, while PIC production diminishes CO2 uptake by 0.27 ± 0.21 Pg C y-1. Without organic carbon production, the SO would be a CO2 source to the atmosphere. Our findings emphasize the importance of DOC and PIC production, in addition to the well-recognized role of POC production, in shaping the influence of carbon export on air-sea CO2 exchange. [ABSTRACT FROM AUTHOR]

Export of sinking particles from the surface ocean is critical for carbon sequestration and to provide energy to the deep biosphere. The magnitude and spatial patterns of this export have been estimated in the past by in situ particle flux observations, satellite‐based algorithms, and ocean biogeochemical models; however, these estimates remain uncertain. Here, we use a recent machine learning reconstruction of global ocean particle size distributions (PSDs) from Underwater Vision Profiler 5 measurements to estimate carbon fluxes by sinking particles (35 μm–5 mm equivalent spherical diameter) from the surface ocean. We combine global maps of PSD properties with empirical relationships constrained against in situ flux observations to calculate particulate carbon export from the euphotic zone (5.8 ± 0.1 Pg C y−1) and annual maximum mixed layer depths (6.1 ± 0.1 Pg C y−1). The new flux reconstructions suggest a less variable seasonal cycle in the tropical ocean and a more persistent export in the Southern Ocean than previously recognized. Smaller particles (less than 418 μm) contribute most of the flux globally, while larger particles become more important at high latitudes and in tropical upwelling regions. Export from the annual maximum mixed layer exceeds that from the euphotic zone over most of the low‐latitude ocean, suggesting shallow particle recycling and net heterotrophy in the deep euphotic zone. These estimates open the way to fully three‐dimensional global reconstructions of particle fluxes in the ocean, supported by the growing database of in situ optical observations. Key Points: A new estimate of sinking particulate carbon fluxes from the surface ocean is developed from reconstructions of particle size distributionSmaller particles contribute more to the total sinking carbon flux than large particlesCarbon flux estimates from two depth horizons suggest net heterotrophy in the deeper euphotic zone, rather than autotrophy [ABSTRACT FROM AUTHOR]

Shipboard sampling of ocean biogeochemical properties is necessarily limited by logistical and practical constraints. As a result, the majority of observations are obtained for the spring/summer period and in regions relatively accessible from a major port. This limitation may bias the conceptual understanding we have of the spatial and seasonal variability in important components of the Earth system. Here we examine the influence of sampling bias on global estimates of carbon export flux by sub-sampling a biogeochemical model to simulate real, realistic and random sampling. We find that both the sparseness and the ‘clumpy’ character of shipboard flux observations generate errors in estimates of globally extrapolated export flux of up to ∼ ± 20%. The use of autonomous technologies, such as the Biogeochemical-Argo network, will reduce the uncertainty in global flux estimates to ∼ ± 3% by both increasing the sample size and reducing clumpiness in the spatial distribution of observations. Nevertheless, determining the climate change-driven trend in global export flux may be hampered due to the uncertainty introduced by interannual variability in sampling patterns.

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.

This contribution provides an introduction to a sequence of five papers (CCE I- CCE V) that describe the impact of the Warm Anomaly of 2014–15 and El Niño 2015–16 on the pelagic food web of the southern California Current Ecosystem. These contributions analyze the influence of these two warm water perturbations on satellite-based measures of ocean fronts, export efficiency out of the euphotic zone, copepod egg production, mesozooplankton community structure, and a synthesis of primary production, mesozooplankton grazing, and gravitational fluxes of organic carbon.

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.

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.

This contribution provides an introduction to a sequence of five papers (CCE I- CCE V) that describe the impact of the Warm Anomaly of 2014–15 and El Niño 2015–16 on the pelagic food web of the southern California Current Ecosystem. These contributions analyze the influence of these two warm water perturbations on satellite-based measures of ocean fronts, export efficiency out of the euphotic zone, copepod egg production, mesozooplankton community structure, and a synthesis of primary production, mesozooplankton grazing, and gravitational fluxes of organic carbon.

This study analyzed the diversity and abundance of diatom frustules including the ancillary parameters using the core top sediments from five locations (21, 19, 15, 13, and 11°N) along the central Arabian Sea (64°E), an area profoundly influenced by atmospheric forcing (monsoons) and oxygen minimum zone (OMZ) with high spatial variability. Significantly higher organic carbon (0.97 ± 0.05%) and diatom frustules (5.92 ± 0.57 × 104 valves g−1) were noticed in the north (21, 19, 15°N) where natural nutrient enrichment via open-ocean upwelling, winter convection, and lateral advection support large diatom-dominated phytoplankton blooms and intense OMZ. Conversely, the south (13, 11°N) depicted significantly lower organic carbon (0.74 ± 0.08%) as well as frustules (4.02 ± 0.87 × 104 valves g−1) as this area mostly remains nutrient-poor dominated by small-medium-sized phytoplankton. The north was dominated by large-sized diatoms like Coscinodiscus that could escape grazing and sink consequently due to higher ballasting. Furthermore, the presence of the intense OMZ in the north might reduce grazing pressure (low zooplankton stock) and mineralization speed facilitating higher phytodetritus transport. Relatively smaller chain-forming centric (Thalassiosira) and pennate diatoms (Pseudo-nitzschia, Fragilaria, Nitzschia, etc.) were found throughout the transect with higher abundance in the south. The euphotic diatom diversity from the existing literature was compared with the frustule diversity from the sediments suggesting not all diatoms make their way to the abyss. Such distinct spatial north–south variability in diatom frustule size as well as abundance could be attributed to cell size, grazing, and water column mineralization rates related to OMZ. [ABSTRACT FROM AUTHOR]

Zooplankton fecal pellets play a significant role in particulate organic carbon (POC) export, which has been widely reported in different ocean regions. However, zooplankton fecal pellet in the tropical marginal sea and its contribution to the total POC flux has not been sufficiently investigated yet. Here, for the first time we report zooplankton fecal pellet characteristics and fluxes in the South China Sea from June 2020 to May 2021 via one mooring with two sediment traps deployed at 500 and 2,190 m depths. Average fecal pellet numerical flux was 2.30 × 104 and 4.22 × 104 pellets m−2 d−1 at 500 and 2,190 m, respectively, corresponding to average fecal pellet carbon (FPC) flux from 0.31 to 0.71 mg C m−2 d−1. Small ellipsoidal and spherical pellets accounted for more than 90% of the total numerical flux, while large cylindrical pellets, although relatively rare (6%), accounted for almost 15% of the FPC flux. Both fecal pellet numerical and carbon fluxes were higher in winter, when the strong East Asian winter monsoon winds and heavy rainfall promoted the marine primary production. Higher fecal pellet fluxes combining with the presence of extra‐large pellets at 2,190 m compared to those at 500 m suggested the in‐situ repackaging of deep‐dwelling zooplankton communities in the mesopelagic and bathypelagic zones. Contribution of fecal pellets to the total POC flux ranged from 0.4% to 30.0% (averaging 9.0%), with higher values occurring during winter monsoon, at which time fecal pellets became a critical conveyor of carbon export to the deep sea. Plain Language Summary: Zooplankton fecal pellets sinking from the sea surface and water column constitute an important component of the marine biological pump. Time series study of fecal pellet flux is therefore of great importance to investigate the deep‐sea particulate organic carbon export. In this study, we examined the temporal and spatial variation of fecal pellet flux in the tropical marginal sea for the first time. Fecal pellet numerical flux and carbon flux at 500 and 2,190 m water depths in the southern South China Sea were quantified, which exhibited great seasonal signals and vertical changes. Strong East Asian winter monsoon contributed to higher fecal pellet fluxes through strengthening the water column mixing and enhancing the marine primary production. In‐situ repackaging of deep‐dwelling zooplankton communities were responsible for the increased fecal pellet flux in the deep sea. Our result of zooplankton fecal pellet flux in the southern South China Sea reflects a dynamic balance between the marine primary production, zooplankton consumption, recycling process, and in‐situ production. Therefore, this study confirms the significant role that zooplankton fecal pellet plays in the local carbon export and provides new insight into the biogeochemical cycling in the tropical marginal sea. Key Points: Zooplankton fecal pellets constitute 0.4%–30.0% of marine particulate organic carbon flux in the deep southern South China SeaStrong East Asian winter monsoon promotes the growth of zooplankton communities and increases their fecal pellet fluxesRepackaging of deep‐dwelling zooplankton communities in mesopelagic and bathypelagic zones contribute to enhanced deep‐sea carbon export [ABSTRACT FROM AUTHOR]

ABSTRACT Picocyanobacteria are the most abundant primary producers in the ocean and play a fundamental role in marine carbon cycling. Quantification of picocyanobacteria on sinking particles and in sediments is essential to understanding their contribution to the biological carbon pump. We designed a primer set targeting the 16S-23S rRNA internal transcribed spacer (ITS) sequence of cyanobacteria and established a quantitative PCR (qPCR) method for quantifying the ITS sequence abundance. High-throughput sequencing confirmed that this primer set can cover broad diversities of marine picocyanobacteria and avoid amplification of other marine cyanobacteria such as Trichodesmium and Crocosphaera. Amplification efficiencies were slightly different when seven marine Synechococcus and Prochlorococcus strains were assayed. The qPCR results were comparable with flow cytometry for water samples. Using this method, we found that, in the dark ocean, picocyanobacterial ITS sequence abundances were 10 to 100 copies/mL in the size fraction of 0.2 to 3 μm, which were 1 to 3 orders of magnitude more abundant than on the >3-μm particles. We also found that picocyanobacterial ITS abundance in sediment ranged from 105 to 107 copies/g along two nearshore-to-offshore transects in the northern South China Sea. These results further explain the important role of picocyanobacteria in carbon export. Collectively, we provide a qPCR method quantifying the total abundance of marine picocyanobacteria on water column particles and in sediments. Moreover, this newly designed primer set can be also applied to investigate the community of picocyanobacteria via high-throughput sequencing. IMPORTANCE Picocyanobacteria are the most abundant primary producers in the ocean. However, quantification of picocyanobacteria on the sinking particles and in sediments remains challenging using flow cytometry or epifluorescence microscopy. Here, we developed a real-time PCR method to quantify picocyanobacteria using a newly designed primer set specifically targeting the 16S-23S rRNA ITS sequence of cyanobacteria. We showed that in the dark ocean, picocyanobacteria are 1 to 3 orders of magnitude more abundant in small particles (0.2 to 3 μm) than in larger particles (>3 μm). This result supports the important role of direct sinking free-living picocyanobacteria cells in the carbon export to deep ocean. We also found that the picocyanobacterial ITS sequence abundance were 105 to 107 copies per gram in sediments, suggesting significant accumulation of sinking picocyanobacteria in the benthic ecosystem. This qPCR method can be used to quantify the contribution of picocyanobacteria to the biological carbon pump.

Although iron and light are understood to regulate the Southern Ocean biological carbon pump, observations have also indicated a possible role for manganese. Low concentrations in Southern Ocean surface waters suggest manganese limitation is possible, but its spatial extent remains poorly constrained and direct manganese limitation of the marine carbon cycle has been neglected by ocean models. Here, using available observations, we develop a new global biogeochemical model and find that phytoplankton in over half of the Southern Ocean cannot attain maximal growth rates because of manganese deficiency. Manganese limitation is most extensive in austral spring and depends on phytoplankton traits related to the size of photosynthetic antennae and the inhibition of manganese uptake by high zinc concentrations in Antarctic waters. Importantly, manganese limitation expands under the increased iron supply of past glacial periods, reducing the response of the biological carbon pump. Overall, these model experiments describe a mosaic of controls on Southern Ocean productivity that emerge from the interplay of light, iron, manganese and zinc, shaping the evolution of Antarctic phytoplankton since the opening of the Drake Passage. Plain Language Summary: Because of the Southern Ocean's unique role in ocean circulation, Antarctic phytoplankton profoundly influence the global carbon cycle. For instance, an increase in the supply of iron—the main nutrient limiting Antarctic phytoplankton—is thought to have lowered CO2 during past ice ages by increasing phytoplankton photosynthesis. However, the potential for other essential elements to limit Southern Ocean productivity is not well known. By accounting for requirements of several nutrients in a global model, we have identified that manganese, an essential cofactor in photosynthesis, can limit phytoplankton growth across the Southern Ocean. The enduring role of manganese deficiency will likely influence the response of Southern Ocean ecosystems to ongoing climate change. Key Points: Mn scarcity in the Southern Ocean limits phytoplankton growth in a global biogeochemical model, especially during austral springThe spatial extent of Mn limitation is sensitive to phytoplankton traits governing photophysiology and metal homeostasisGreater dust deposition to the Southern Ocean expands the role of Mn limitation and restricts carbon export from Fe fertilization [ABSTRACT FROM AUTHOR]