Photosynthesis in the surface ocean produces approximately 100 gigatonnes of organic carbon per year, of which 5 to 15 per cent is exported to the deep ocean. The rate at which the sinking carbon is ...converted into carbon dioxide by heterotrophic organisms at depth is important in controlling oceanic carbon storage. It remains uncertain, however, to what extent surface ocean carbon supply meets the demand of water-column biota; the discrepancy between known carbon sources and sinks is as much as two orders of magnitude. Here we present field measurements, respiration rate estimates and a steady-state model that allow us to balance carbon sources and sinks to within observational uncertainties at the Porcupine Abyssal Plain site in the eastern North Atlantic Ocean. We find that prokaryotes are responsible for 70 to 92 per cent of the estimated remineralization in the twilight zone (depths of 50 to 1,000 metres) despite the fact that much of the organic carbon is exported in the form of large, fast-sinking particles accessible to larger zooplankton. We suggest that this occurs because zooplankton fragment and ingest half of the fast-sinking particles, of which more than 30 per cent may be released as suspended and slowly sinking matter, stimulating the deep-ocean microbial loop. The synergy between microbes and zooplankton in the twilight zone is important to our understanding of the processes controlling the oceanic carbon sink.
Atmospheric carbon dioxide levels are strongly controlled by the depth at which the organic matter that sinks out of the surface ocean is remineralized. This depth is generally estimated from ...particle flux profiles measured using sediment traps. Inherent in this analysis is a steady state assumption that export from the surface does not significantly change in the time it takes material to reach the deepest trap. However, recent observations suggest that a significant fraction of material in the mesopelagic zone sinks slowly enough to bring this into doubt. We use data from a study in the North Atlantic during July/August 2009 to challenge the steady state assumption. An increase in biogenic silica flux with depth was observed which we interpret, based on vertical profiles of diatom taxonomy, as representing the remnants of the spring diatom bloom sinking slowly (<40 m d−1). We were able to reproduce this behavior using a simple model using satellite‐derived export rates and literature‐derived remineralization rates. We further provide a simple equation to estimate “additional” (or “excess”) particulate organic carbon supply to the dark ocean during nonsteady state conditions, which is not captured by traditional sediment trap deployments. In seasonal systems, mesopelagic net organic carbon supply could be wrong by as much as 25% when assuming steady state. We conclude that the steady state assumption leads to misinterpretation of particle flux profiles when input fluxes from the upper ocean vary on the order of weeks, such as in temperate and polar regions with strong seasonal cycles in export.
Key Points
Increased biogenic silica fluxes with depth imply nonsteady state conditions in the NE Atlantic in August 2009
Taxonomic flux analysis and simple modeling support the nonsteady state hypothesis
By assuming steady state, net organic carbon supply to the dark ocean may be wrong by ≤ 25%
Sinking organic particles transfer ∼10 gigatonnes of carbon into the deep ocean each year, keeping the atmospheric CO₂ concentration significantly lower than would otherwise be the case. The exact ...size of this effect is strongly influenced by biological activity in the ocean's twilight zone (∼50–1,000 m beneath the surface). Recent work suggests that the resident zooplankton fragment, rather than ingest, the majority of encountered organic particles, thereby stimulating bacterial proliferation and the deep‐ocean microbial food web. Here we speculate that this apparently counterintuitive behaviour is an example of ‘microbial gardening’, a strategy that exploits the enzymatic and biosynthetic capabilities of microorganisms to facilitate the ‘gardener's’ access to a suite of otherwise unavailable compounds that are essential for metazoan life. We demonstrate the potential gains that zooplankton stand to make from microbial gardening using a simple steady state model, and we suggest avenues for future research.
Abstract
Recognition of the importance of jellyfish in marine ecosystems is growing. Yet, the biochemical composition of the mucus that jellyfish constantly excrete is poorly characterized. Here we ...analyzed the macromolecular (proteins, lipids and carbohydrates) and elemental (carbon and nitrogen) composition of the body and mucus of five scyphozoan jellyfish species (Aurelia aurita, Chrysaora fulgida, Chrysaora pacifica, Eupilema inexpectata and Rhizostoma pulmo). We found that the relative contribution of the different macromolecules and elements in the jellyfish body and mucus was similar across all species, with protein being the major component in all samples (81 ± 4% of macromolecules; 3.6 ± 3.1% of dry weight, DW) followed by lipids (13 ± 4% of macromolecules; 0.5 ± 0.4%DW) and carbohydrates (6 ± 3% of macromolecules; 0.3 ± 0.4%DW). The energy content of the jellyfish matter ranged from 0.2 to 3.1 KJ g−1 DW. Carbon and nitrogen content was 3.7 ± 3.0 and 1.0 ± 0.8%DW, respectively. The average ratios of protein:lipid:carbohydrate and carbon:nitrogen for all samples were 14.6:2.3:1 and 3.8:1, respectively. Our study highlights the biochemical similarity between the jellyfish body and mucus and provides convenient and valuable ratios to support the integration of jellyfish into trophic and biogeochemical models.
The remineralization depth of particulate organic carbon (POC) fluxes exported from the surface ocean exerts a major control over atmospheric CO₂ levels. According to a long‐held paradigm most of the ...POC exported to depth is associated with large particles. However, recent lines of evidence suggest that slow‐sinking POC (SSPOC) may be an important contributor to this flux. Here we assess the circumstances under which this occurs. Our study uses samples collected using the Marine Snow Catcher throughout the Atlantic Ocean, from high latitudes to midlatitudes. We find median SSPOC concentrations of 5.5 μg L−1, 13 times smaller than suspended POC concentrations and 75 times higher than median fast‐sinking POC (FSPOC) concentrations (0.07 μg L−1). Export fluxes of SSPOC generally exceed FSPOC flux, with the exception being during a spring bloom sampled in the Southern Ocean. In the Southern Ocean SSPOC fluxes often increase with depth relative to FSPOC flux, likely due to midwater fragmentation of FSPOC, a process which may contribute to shallow mineralization of POC and hence to reduced carbon storage. Biogeochemical models do not generally reproduce this behavior, meaning that they likely overestimate long‐term ocean carbon storage.
Key Points
Suspended, slow‐, and fast‐sinking particulate organic carbon (POC) contributes 94:6:<1% to total POC concentration, respectively
POC flux below the mixed layer is often dominated by slow‐sinking particles
In situ generation of slow‐sinking POC below the mixed layer, via midwater fragmentation, may lead to shallower mineralization of particles
Prokaryotes play a central role in aquatic ecosystems by consuming approximately half of the organic matter produced by aquatic primary production, of which a fraction is used for growth. Accurately ...measuring this prokaryotic biomass production is key to understanding aquatic carbon and nutrient cycles, since it is instrumental in driving biogeochemical processes that control parameters such as atmospheric carbon content. Aquatic prokaryotic biomass production is typically estimated from incorporation rates of the amino acid leucine during radiotracer experiments—a method widely used since the 1980s. Here we evaluate the underlying assumptions of the method with a focus on the associated conversion factors and review them in the context of empirical data. We demonstrate that the commonly used theoretical conversion factors fail to account for leucine's use as precursor for de novo protein synthesis and its respiration. As a consequence, prokaryotic biomass production is likely considerably overestimated when applying the standard conversion factors. Most severely affected are open‐ocean, mesopelagic and benthic environments, where 25% of the estimates are likely to be overestimated by at least a factor of 6.1, 4.9, and 6.5, respectively. We propose a refined carbon‐to‐leucine conversion factor and make recommendations for improving and selecting appropriate experimental protocols.
In situ imaging of particles in the ocean are rapidly establishing themselves as powerful tools to investigate the ocean carbon cycle, including the role of sinking particles for carbon sequestration ...via the biological carbon pump. A big challenge when analysing particles in camera images is determining the size of the particle, which is required to calculate carbon content, sinking velocity and flux. A key image processing decision is the algorithm used to decide which part of the image forms the particle and which is the background. However, this critical analysis step is often unmentioned and its effect rarely explored. Here we show that final flux estimates can easily vary by an order of magnitude when selecting different algorithms for a single dataset. We applied a range of static threshold values and 11 different algorithms (7 threshold and 4 edge detection algorithms) to particle profiles collected by the LISST-Holo system in two contrasting environments. Our results demonstrate that the particle detection method does not only affect estimated particle size but also particle shape. Uncertainties are likely exacerbated when different particle detection methods are mixed, e.g., when datasets from different studies or devices are merged. We conclude that there is a clear need for more transparent method descriptions and justification for particle detection algorithms, as well as for a calibration standard that allows intercomparison between different devices.
The Agulhas Bank on the tip of southern Africa, like other shelf seas, is a relatively productive environment which plays a crucial role in the biology and success of many commercially valuable fish ...species. Fish and their larvae depend on zooplankton to feed on but, despite their importance, little is known about zooplankton distribution and production on the Agulhas Bank. Here we present results from a survey conducted in March 2019 on the East and Central Agulhas Bank, investigating mesozooplankton abundance, biovolume, taxonomic composition, size distribution (normalised biovolume size spectrum (NBSS) approach) and secondary production. A clear cross-shore gradient was observed with the inner-shelf having higher abundance and biovolume of mesozooplankton dominated by small-size organisms, most likely mirroring higher overall productivity of the coastal waters, while the outer-shelf showed the opposite trend (i.e., low abundance and biovolume; shallow NBSS slopes). This general pattern on the outer-shelf was, however, disturbed in one location (between 24 and 25°E) with a distinguishable mesozooplankton community, most likely linked to the passage of a meander on the inshore side of the Agulhas Current. The Central Agulhas Bank was typified by high mesozooplankton biomass (∼4 g C m-2), comparable to upwelling areas and dominated by copepods and doliolids. High copepod biomass was observed in this region before and was linked to a feature called the “Cold Ridge”. However, during our survey, no such ridge was observed. The mixed layer depth was relatively deep (>20 m) and high Chlorophyll a concentration was measured at depth, despite low net primary production rates. We suggest that this region is prone to other mechanisms such as retention due to cyclonic circulation or processes injecting nutrients in the upper mixed layer that require further research. Secondary production on the Agulhas Bank was in the same range as other shelf seas (0.03–1.55 g C m-2 d-1) and was correlated with mesozooplankton biomass. The comparison of primary and secondary production, measured simultaneously, suggested that mesozooplankton exert a significant control on net primary production, and can, in some areas, be food-limited (e.g., Central Agulhas Bank and inshore waters).
•Mesozooplankton biomass on the Agulhas Bank in March was 0.1–4.27 g C m-2.•The inner-shelf reveals higher zooplankton biomass and steeper NBSS slopes.•The Central Agulhas Bank has high zooplankton biomass, comparable to upwelling areas.•Secondary production in March ranged from 0.03 to 1.55 g C m-2 d-1.
Vertical particle fluxes of particulate organic carbon (POC), chlorophyll a (Chl a) and biogenic silica (bSi) were measured on the productive shelf of southern Africa, the Agulhas Bank (AB), in March ...2019. Sinking particulate material in the form of aggregates is hypothesized to form the benthic nepheloid layer (BNL) which is a turbid layer found near the seabed. This layer is known to affect the spawning success of squid as it is linked to high turbidity which reduces visibility during mating. To determine the distribution of fluxes and particle composition in the AB, we collected water samples below the surface mixed layer (‘export’) and near the seabed (‘bottom’) using a Marine Snow Catcher. POC export fluxes were significantly higher inshore than offshore (mean ± SD: 944.6 ± 302.0 & 461.1 ± 162.1 mg POC m−2 d−1, respectively). There was no significant difference in the cross-shelf distribution of Chl a and bSi export fluxes, however the inshore fluxes of Chl a and bSi were higher than offshore, suggesting a link between export fluxes and sinking organic matter derived from the more productive inshore surface waters. All bottom fluxes were significantly higher inshore, suggesting the contribution of sinking organic particles and resuspended bottom sediments to inshore fluxes. POC export efficiency (ratio of exported POC flux relative to net primary production (NPP)) was higher on the AB (range: 0.58–9.56) compared to the global shelf seas ratio of 0.18 and not related to NPP, suggesting an export of standing stock of carbon biomass, likely produced before the cruise. Transfer efficiency (i.e., the amount of exported flux that reaches the bottom) was also high (max: 0.99, 1.0 and 33.04 for POC, Chl a and bSi, respectively) but did not show a clear spatial pattern. We observed a significant positive correlation between bottom turbidity (a proxy for BNL presence) and export POC flux, suggesting the possibility that sinking organic matter is contributing to BNL formation on the AB.
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