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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]

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]

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]

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.

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]

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 (

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.

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.

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.

Climate change is expected to alter the intensity and frequency of upwelling in high productive coastal regions, thus impacting nutrient fluxes, primary productivity and consequently carbon cycling. However, it is unknown how these changes will impact the planktonic (phytoplankton and bacteria) community structure, which affects community respiration (CR) and hence the carbon available for sequestration or transfer to upper trophic levels. Here we present results from a 37-day mesocosm experiment where we examined the response of CR to nutrient additions by simulating upwelling events at different intensities (low, medium, high and extreme) and modes (singular and recurring additions). We also analysed the potential contribution of different plankton size classes and functional groups to CR. The trend in accumulated CR with respect to nutrient fertilisation (total nitrogen added during the experiment) was linear in the two modes. Microplankton (mostly diatoms) and nanoplankton (small flagellates) dominated under extreme upwelling intensities and high CR in both singular and recurring upwelling modes, explaining >65% of the observed variability in CR. In contrast, prokaryotic picoplankton (heterotrophic bacteria and autotrophic cyanobacteria) explained

A high resolution coastal and ocean hydrodynamic model system was used to investigate the transport and deposition patterns of Particulate Organic Matter (POM) from kelp farmed at three locations of different properties: a sheltered location, an exposed location, and an offshore location. Published values on the sinking speeds of organic particles from kelp were used, spanning several orders of magnitude. Recent work on quantifying the release of particulate organic matter from farmed kelp was used to link the release of carbon to possible cultivation volumes and scenarios, and finally to link this to the potential for carbon loading on the ocean floor. The results are presented in terms of loading and distribution per unit harvested kelp, and the loading estimates are compared with estimates of natural (background) primary production. According to the simulation results, organic matter may be transported anything from a few (hundred) meters up to a hundred km away from the release site, depending on the sinking rates, time of release, and the location. The depth at which the matter settles on the sea floor likewise depends on the properties of the matter and the sites. The time until settlement varied from minutes to several hundred hours. The results underscore the importance of constraining the dispersal and deposition of detritus from kelp cultivation in order to better understand and quantify associated environmental risks posed by organic loading, and the potential for seafloor carbon sequestration by kelp farming as a nature based climate solution.

A freshwater-induced barrier layer (BL) is a common physical phenomenon both in coastal waters and the open ocean. To examine the effects of BL on the biological production and the associated carbon export, a physical-biogeochemical survey was conducted in the Bay of Bengal. Severe depletions of surface phosphorus and the deepening of the nutricline were observed at the BL-affected stations due to the vertical mixing prohibition. The lowered surface chlorophyll a (Chl a) and squeezed deep Chl a maximum (DCM) layer also resulted in the ~18% lowered vertically integrated Chl a at the said stations. The composition of the net-sampled zooplankton was altered, and the abundance decreased by half at the BL-affected station (29.68 ind. m−3) compared with the unaffected station (55.52 ind. m−3). Such reductions in major zooplankton groups were confirmed by a video plankton recorder (VPR). The VPR observation indicated that there was a lower (by one-half) abundance of detritus at the BL-affected station, while the much lower carbon export flux rates were estimated to be at the BL-affected station (0.31 mg C m−2 d−1) rather than the unaffected station (0.77 mg C m−2 d−1). An idealized one-dimensional nutrient-phytoplankton-detritus model identified that the existence of BL can lead to decreased surface nutrients and phytoplankton concentrations, squeezed DCM layers, and lower detritus abundances. Finally, this study indicated that BL layers inhibit biological production and reduce carbon export, thus playing an important role in the ocean biogeochemical cycles.

The net primary productivity (NPP) of marine phytoplankton is ∼50 Pg C year–1, and roughly 10–20% of this NPP is exported out of the surface ocean as sinking particulate organic carbon (POC). Numerous mechanisms are hypothesized to control POC export out of the surface ocean but the relative importance of the various mechanisms remains poorly quantified on global scales. Here, we use a previously published satellite-based mechanistic model of POC export to examine the effects on global POC export of size-specific physical aggregation, size-specific and temperature-dependent zooplankton fecal pellet production, and size-specific and temperature-dependent non-grazing phytoplankton mortality. We test these mechanisms in different model configurations to determine if these processes improve the ability of the model to match POC export observations, and to assess the role of each process in controlling global POC export. We find that all model configurations predict that over 60% of the global POC export is from small zooplankton fecal pellets. All model configurations predict similar total POC export, and we find only small differences in the magnitude, timing, and geographical variations of total POC export. However, the fraction of total POC export due to sinking phytoplankton aggregates, and that due to the fecal pellets of large zooplankton, vary by more than a factor of two across the different model configurations. The POC export in all models is most sensitive to parameters controlling zooplankton fecal fluxes and non-grazing phytoplankton mortality. We compared zooplankton grazing rates predicted by the models to results of experimental data, and found that some models match the experimental grazing rates better than others, although data uncertainties remain large. More field measurements of bulk ecosystem rates (i.e., phytoplankton aggregation and zooplankton grazing), as well as explicit determinations of of the proportion of fecal matter to phytoplankton aggregation, will help to better constrain mechanistic models of global POC export.

Optical instruments can rapidly determine numbers and characteristics of water column particles with high sensitivity. Here we show the usefulness of optically assessed total particle volume below the main pycnocline to estimate carbon export in two systems: the open subarctic North Atlantic and the Ross Sea, Antarctica. Both regions exhibit seasonally high phytoplankton production and efficient export (i.e., a strong biological pump). Total particle volumes in the mesopelagic (200–300 m) were significantly correlated with those in the overlying surface mixed layer (50–60 m), indicating that most particles at depth reflect export from the surface. This connectivity, however, is modulated by the physical structure of the water column and by particle type (e.g., the presence of colonies of the haptophyte Phaeocystis antarctica versus diatoms). Evidence from both regions show that a strong pycnocline can delay or may even prevent particles from settling to deeper layers, which then succumb to disintegration, and microbial and zooplankton consumption. Strong katabatic winds in the Ross Sea may deepen the mixed layer, causing a rapid transfer of particles to mesopelagic depths through the mixed-layer pump. Independent estimates of seasonally integrated export production in the Ross Sea, based on upper water column carbon mass balance, were significantly correlated (in the order of shared variance) with (1) total particle volumes from images, (2) particulate organic carbon, and (3) chlorophyll fluorescence, all recorded at a depth range of 200–300 m. Carbon export was not significantly correlated with particle abundance measured by a Coulter counter at the same depth range. Measuring total particle volume below the primary pycnocline is therefore a useful approach to estimate carbon export at least in regions characterized by seasonally high particle export.

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 = ∑i=1xni⁢A⁢diB⁢Δ⁢di), 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.

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.

The depth-attenuation of sinking particulate organic carbon (POC) is of particular importance for the ocean's role in the global carbon cycle. Numerous idealized flux-vs.-depth relationships are available to parameterize this process in Earth System Models. Here we show that these relationships are statistically indistinguishable from available POC flux profile data. Despite their quantitative similarity, we also show these relationships have very different implications for the flux leaving the upper ocean, as well as for the mechanisms governing POC flux. We discuss how this tension might be addressed both observationally and in modeling studies.

Estimates of dissolved organic carbon (DOC) release by marine macrophyte communities (seagrass meadows and macroalgal beds) based on in situ benthic chambers from published and unpublished are compiled in this study. The effect of temperature and light availability on DOC release by macrophyte communities was examined. Almost 85 % of the seagrass communities and all of macroalgal communities examined acted as net sources of DOC. Net DOC fluxes in seagrass communities increase positively with water temperature. In macroalgal communities net DOC fluxes under light exceeded those under dark condition, however, this trend was weaker in seagrass communities. Shading of a mixed seagrass meadow in The Philippines led to a significant reduction on the net DOC release when shading was maintained for 6 days compared to only 2 days of shading. Net DOC fluxes increased with increasing community respiration, but were independent of primary production or net community production. The estimated global net DOC flux, and hence export, from marine macrophytes is about 0.158 ± 0.055 Pg C yr-1 or 0.175 ± 0.056 Pg C yr-1 depending on the global extent of seagrass meadows considered.