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1.
The Lagrangian Southern Ocean Iron Release Experiment (SOIREE) allowed study of a gradually evolving iron-mediated phytoplankton bloom in water labelled with the inert tracer sulfur hexafluoride, SF6. This article describes a pelagic carbon budget for the mixed layer in SOIREE and assesses the extent to which closure of the budget is achieved. Net community production (NCP) converted 837 mmol m−2 of inorganic carbon to organic carbon in 12.0 d after the first iron addition. A large fraction (41%) of NCP remained as particulate organic carbon in the mixed layer of the iron-enriched patch, while 23% was lost by horizontal dispersion and 0–29% was exported. The closure of the carbon budget is hampered by the lack of measurements of dissolved organic carbon (DOC), by a major uncertainty in carbon export, and by use of empirical conversion factors in estimates of carbon biomass and metabolic rates. Lagrangian carbon-budget studies may be improved by direct measurement of all major carbon parameters and conversion factors. Carbon cycling in the SOIREE bloom resembled that in ‘natural’ algal blooms in the open Southern Ocean in some respects, but not in all. Daily NCP in the SOIREE bloom (70 mmol m−2 d−1) was higher than in natural blooms, partly because other studies did not account for horizontal dispersion, were for longer periods or included less productive areas. The build-up of POC stock and carbon export as a fraction of NCP in SOIREE were in the lower range of observations elsewhere.  相似文献   

2.
Young Sound is a deep-sill fjord in NE Greenland (74°N). Sea ice usually begins to form in late September and gains a thickness of 1.5 m topped with 0–40 cm of snow before breaking up in mid-July the following year. Primary production starts in spring when sea ice algae begin to flourish at the ice–water interface. Most biomass accumulation occurs in the lower parts of the sea ice, but sea ice algae are observed throughout the sea ice matrix. However, sea ice algal primary production in the fjord is low and often contributes only a few percent of the annual phytoplankton production. Following the break-up of ice, the immediate increase in light penetration to the water column causes a steep increase in pelagic primary production. Usually, the bloom lasts until August–September when nutrients begin to limit production in surface waters and sea ice starts to form. The grazer community, dominated by copepods, soon takes advantage of the increased phytoplankton production, and on an annual basis their carbon demand (7–11 g C m−2) is similar to phytoplankton production (6–10 g C m−2). Furthermore, the carbon demand of pelagic bacteria amounts to 7–12 g C m−2 yr−1. Thus, the carbon demand of the heterotrophic plankton is approximately twice the estimated pelagic primary production, illustrating the importance of advected carbon from the Greenland Sea and from land in fuelling the ecosystem.In the shallow parts of the fjord (<40 m) benthic primary producers dominate primary production. As a minimum estimate, a total of 41 g C m−2 yr−1 is fixed by primary production, of which phytoplankton contributes 15%, sea ice algae <1%, benthic macrophytes 62% and benthic microphytes 22%. A high and diverse benthic infauna dominated by polychaetes and bivalves exists in these shallow-water sediments (<40 m), which are colonized by benthic primary producers and in direct contact with the pelagic phytoplankton bloom. The annual benthic mineralization is 32 g C m−2 yr−1 of which megafauna accounts for 17%. In deeper waters benthic mineralization is 40% lower than in shallow waters and megafauna, primarily brittle stars, accounts for 27% of the benthic mineralization. The carbon that escapes degradation is permanently accumulated in the sediment, and for the locality investigated a rate of 7 g C m−2 yr−1 was determined.A group of walruses (up to 50 adult males) feed in the area in shallow waters (<40 m) during the short, productive, ice-free period, and they have been shown to be able to consume <3% of the standing stock of bivalves (Hiatella arctica, Mya truncata and Serripes Groenlandicus), or half of the annual bivalve somatic production. Feeding at greater depths is negligible in comparison with their feeding in the bivalve-rich shallow waters.  相似文献   

3.
Mesoscale eddies may enhance primary production (PP) in the open ocean by bringing nutrient-rich deep waters into the euphotic zone, potentially leading to increased transport of particles to depth. This hypothesis remains controversial, however, due to a paucity of direct particle export measurements. In this study, we investigated particle dynamics using 234Th–238U disequilibria within a mesoscale cold-core eddy, Cyclone Opal, which formed in the lee of the Hawaiian Islands. 234Th samples were collected along two transects across Cyclone Opal as well as during a time-series within the eddy core during a decaying diatom bloom. Particulate carbon (PC), particulate nitrogen (PN) and biogenic silica (bSiO2) fluxes at 150 m varied spatially and temporally within the eddy and strongly depended on the 234Th model formulation used (e.g., steady state versus non-steady state, inclusion of upwelling, etc.). Particle fluxes estimated from a steady state model assuming an upwelling rate of 2 m day−1 yielded the best fit to sediment-trap data. These 234Th-derived particle fluxes ranged from 332±14 to 1719±53 μmol C m−2 day−1, 27±3 to 114±12 μmol N m−2 day−1, and 33±20 to 309±73 μmol Si m−2 day−1. Although PP rates within Cyclone Opal were elevated by a factor of 2–3, PC and PN fluxes were the same, within error, inside and outside of Cyclone Opal. The ratio of PC export to PP remained surprisingly low at <0.03 and similar to those measured in surrounding waters. In contrast, bSiO2 fluxes within the eddy core were three times higher. Detailed analyses of 234Th depth profiles consistently showed excess 234Th at 100–175 m, associated with the remineralization and possible accumulation of suspended and dissolved organic matter from the surface. We suggest that strong microzooplankton grazing facilitated particulate organic matter recycling and resulted in the export of empty diatom frustules. Thus, while eddies may increase PP, they do not necessarily increase PC and PN export to deep waters. This may be a general characteristic of wind-driven cyclonic eddies of the North Pacific Subtropical Gyre and suggests that eddies may preferentially act as a silica pump, thereby playing an important role in promoting silicic-acid limitation in the region.  相似文献   

4.
The Wadden Sea (North Sea, Europe) is a shallow coastal sea with high benthic and pelagic primary production rates. To date, no studies have been carried out in the Wadden Sea that were specifically designed to study the relation between pelagic respiration and production by comparable methods. Because previous studies have suggested that the import of primary-produced pelagic organic matter is important for benthic Wadden Sea carbon budgets, we hypothesised that on an annual average the northern Wadden Sea water column is autotrophic. To test this hypothesis, we studied annual dynamics of primary production and respiration at a pelagic station in a shallow tidal basin (List Tidal Basin, northern Wadden Sea). Since water depth strongly influences production estimates, we calculated primary production rates per unit area in two ways: on the basis of the mean water depth (2.7 m) and on the basis of 1 m depth intervals and their respective spatial extent in the List Tidal Basin. The latter more precise estimate yielded an annual primary production of 146 g C m− 2 y− 1. Estimates based on the mean water depth resulted in a 40% higher annual rate of 204 g C m− 2 y− 1. The total annual pelagic respiration was 50 g C m− 2 y− 1. The P/R ratio varied between seasons: from February to October the water column was autotrophic, with the highest P/R ratio of 4–5 during the diatom spring bloom in April/May. In autumn and winter the water column was heterotrophic. On an annual average, the water column of the List Tidal Basin was autotrophic (P/R 3). We suggest that a large fraction of the pelagic produced organic matter was respired locally in the sediment.  相似文献   

5.
As part of E-Flux III cruise studies in March 2005, we investigated phytoplankton community dynamics in a cyclonic cold-core eddy (Cyclone Opal) in the lee of the Hawaiian Islands. Experimental incubations were conducted under in situ temperature and light conditions on a drift array using a two-treatment dilution technique. Taxon-specific estimates of growth, grazing and production rates were obtained from analyses of incubation results based on phytoplankton pigments, flow cytometry and microscopy. Cyclone Opal was sampled at a biologically and physically mature state, with an 80–100 m doming of isopycnal surfaces in its central region and a deep biomass maximum of large diatoms. Depth-profile experimentation defined three main zones. The upper (mixed) zone (0–40 m), showed little compositional or biomass response to eddy nutrient enrichment, but growth, grazing and production rates were significantly enhanced in this layer relative to the ambient community outside of the eddy. Prochlorococcus spp. dominated the upper mixed layer, accounting for 50–60% of its estimated primary production both inside and outside of Opal. In contrast, the deep zone of 70–90 m showed little evidence of growth rate enhancement and was principally defined by a 100-fold increase of large (>20-μm) diatoms and a shift from Prochlorococcus to diatom dominance (80%) of production. The intermediate layer of 50–60 m marked the transition between the upper and lower extremes but also contained an elevated biomass of physiologically unhealthy diatoms with significantly depressed growth rates and proportionately greater grazing losses relative to diatoms above or below. Microzooplankton grazers consumed 58%, 65% and 55%, respectively, of the production of diatoms, Prochlorococcus and the total phytoplankton community in Cyclone Opal. The substantial grazing impact on diatoms suggests that efficient recycling was the major primary fate of diatom organic production, consistent with the low export fluxes and selective export of biogenic silica, as empty diatom frustules, in Cyclone Opal.  相似文献   

6.
The least known component of the “biological pump” is the active transport of carbon and nutrients by diel vertical migration of zooplankton. We measured CO2 respiration and dissolved organic carbon (DOC) excretion by individual species of common vertically migrating zooplankton at the US JGOFS Bermuda Atlantic Time-series Study (BATS) station. The inclusion of DOC excretion in this study builds on published research on active transport by respiration of inorganic carbon and allows a direct assessment of the role of zooplankton in the production of dissolved organic matter used in midwater microbial processes. On average, excretion of DOC makes up 24% (range=5–42%) of the total C metabolized (excreted+respired) and could represent a significant augmentation to the vertical flux that has already been documented for respiratory CO2 flux by migrant zooplankton. Migratory fluxes were compared to other transport processes at BATS. Estimates of combined active transport of CO2 and DOC by migrators at BATS averaged 7.8% and reached 38.6% of mean sinking POC flux at 150 m, and reached 71.4% of mean sinking POC flux at 300 m. DOC export by migrator excretion averaged 1.9% and reached 13.3% of annual DOC export by physical mixing at this site. During most of the year when deep mixing does not occur, diel migration by zooplankton could provide a supply of DOC to the deeper layers that is available for use by the microbial community. A carbon budget comparing migrant zooplankton transport to the balance of fluxes in the 300–600 m depth strata at BATS shows on average that the total migrant flux supplies 37% of the organic carbon remineralized in this layer, and that migrant DOC flux is more than 3 times the DOC flux gradient by diapycnal mixing. New estimates of active transport of both organic and inorganic carbon by migrants may help resolve observed imbalances in the C budget at BATS, but the magnitude is highly dependent on the biomass of the migrating community.  相似文献   

7.
We used a new experimental device called PASS (PArticle Sinking Simulator) during MedFlux to simulate changes in in situ hydrostatic pressure that particles experience sinking from mesopelagic to bathypelagic depths. Particles, largely fecal pellets, were collected at 200 m using a settling velocity NetTrap (SV NetTrap) in Ligurian Sea in April 2006 and incubated in high-pressure bottles (HPBs) of the PASS system under both atmospheric and continuously increasing pressure conditions, simulating the pressure change experienced at a sinking rate of 200 m d−1. Chemical changes over time were evaluated by measuring particulate organic carbon (POC), carbohydrates, transparent exopolymer particles (TEP), amino acids, lipids, and chloropigments, as well as dissolved organic carbon (DOC) and dissolved carbohydrates. Microbial changes were evaluated microscopically, using diamidinophenylindole (DAPI) stain for total cell counts and catalyzed reporter deposition-fluorescence in situ hybridization (CARD-FISH) for phylogenetic distinctions. Concentrations (normalized to POC) of particulate chloropigments, carbohydrates and TEP decreased under both sets of incubation conditions, although less under the increasing pressure regime than under atmospheric conditions. By contrast, dissolved carbohydrates (normalized to DOC) were higher after incubation and significantly higher under atmospheric conditions, suggesting they were produced at the expense of the particulate fraction. POC-normalized particulate wax/steryl esters increased only under pressure, suggesting biochemical responses of prokaryotes to the increasing pressure regime. The prokaryotic community initially consisted of 43% Bacteria, 12% Crenarchaea and 11% Euryarchaea. After incubation, Bacteria dominated (90%) the prokaryote community in all cases, with γ-Proteobacteria comprising the greatest fraction, followed by the Cytophaga–Flavobacter cluster and α-Proteobacteria group. Using the PASS system, we obtained chemical and microbial evidence that degradation by prokaryotes associated with fecal pellets sinking through mesopelagic waters is limited by the increasing pressure they experience.  相似文献   

8.
《Marine Chemistry》2007,103(1-2):61-75
The capacity of natural dissolved organic matter (DOM) produced during the onset, development and decay of a simulated bloom of the marine diatom Skeletonema costatum to complex free copper has been followed for a 2 week period. Copper binding capacity of the culture was measured by anodic stripping voltammetry (ASV) with a hanging mercury drop electrode (HMDE). The concentration of dissolved organic carbon (DOC) and two fluorophores, M (humic-like, Ex/Em: 320 nm/410 nm) and T (protein-like, Ex/Em: 280 nm/350 nm), were followed during the course of the incubation. Models using DOC concentrations alone could not accurately predict the complexing capacity of the culture, especially at the end of the bloom, and better predictions were obtained when fluorescence measurements were considered. They were helpful in characterising two types of copper ligands produced in the culture. The first type, traced by the fluorescence of peak T, was related to labile DOC directly exuded by phytoplankton. The second type, traced by the fluorescence of peak M, was the refractory humic-like material presumably produced in situ as a by-product of the bacterial degradation of phytogenic materials. During the onset and development of the bloom (days 0 to 7), the fluorescence of peak T explains 60–80% of the total complexing capacity of the culture, suggesting that exuded “protein-like” compounds among other exuded complexing agents efficiently complexed free copper. On the contrary, during the decay (days 8 to 13), these ligands were replaced by humic substances as the complexing agent for copper.  相似文献   

9.
Diatoms play a significant role in the global carbon cycle through their role in biogenic silica production and the transport of organic matter to the seafloor. Recent work has shown that silicified diatom frustules contain a significant amount of organic matter, and that the proportion of diatom-bound organic matter increases with depth in the water column and sediments. Here, we investigate the association between organic matter and the mineral phase. We used a combination of scanning transmission X-ray microscopy (STXM) and carbon X-ray absorption near-edge structure (XANES) spectroscopy to characterize the distribution and composition of organic matter in frustules of the diatom Cylindrotheca closterium and a biomimetic silica gel. To our knowledge, this study represents the first successful attempt to simultaneously image and obtain chemical information about the organic matter within a diatom frustule using X-ray spectromicroscopy near the carbon edge. Organic carbon, most likely protein, was distributed throughout the frustules and was not removed by harsh chemical treatment. The physical structure of the frustules appeared to be related to the chemical composition of this organic matter, with aromatic or unsaturated carbon being concentrated in the most intricately patterned regions of the frustule. A similar physical and chemical structure was observed in a biomimetic silica gel precipitated spontaneously with polylysine. These results are consistent with the theory that organic constituents of diatom frustules direct silica precipitation and become incorporated within the silica matrix as it forms. The relationship between organic matter composition and silica morphology, the failure of harsh chemical treatments to remove this organic matter, and the spontaneous nature of the co-precipitation of silica and organic matter indicate some chemical interaction between the siliceous and organic components of diatom frustules. Frustule-bound organic matter should therefore be protected from decomposition in the water column or diagenetic alteration in sediments unless the frustule dissolves.  相似文献   

10.
Two processes are generally explained as causes of temporal changes in the stoichiometric silicon/nitrogen (Si/N) ratios of sinking particles and of nutrient consumption in the surface water during the spring diatom bloom: (1) physiological changes of diatom under the stress of photosynthesis of diatom and (2) differences of regeneration between silicon and nitrogen. We investigated which process plays an important role in these changes using a one-dimensional ecosystem model that explicitly represents diatom and the other non-silicious phytoplankton. The model was applied to station A7 (41°30′ N, 145°30′ E) in the western North Pacific, where diatom regularly blooms in spring. Model simulations show that the Si/N ratios of the flux exported by the sinking particles at 100 m depth and of nutrient consumptions in the upper 100 m surface water have their maxima at the end of the spring diatom bloom, the values and timings of which are significantly different from each other. Analyses of the model results show that the differences of regeneration between silicon and nitrogen mainly cause the temporal changes of the Si/N ratios. On the other hand, the physiological changes of diatoms under stress can hardly cause these temporal changes, because the effect of the change in the diatom's uptake ratio of silicon to nitrogen is cancelled by that in its sinking rate.  相似文献   

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