Biological components of Ross Sea short-term particle fluxes in the austral summer of 1995–1996

The Ross Sea, a region of high seasonal production in the Southern Ocean, is characterized by blooms of the haptophyte Phaeocystis antarctica and of diatoms. The different morphology, structural composition and consumption of these two phytoplankton by grazing zooplankton may result in different carbon cycling dynamics and carbon flux from the euphotic zone. We sampled short-term (2 days) particle flux at 5 sites from 177.6°W to 165°E along a transect at 76.5°S with traps placed below the euphotic zone at 200 m during December 1995–January 1996. We estimated carbon flux of as many eucaryotic organisms and fecal pellets as possible using microscopy for counts and measurements and applying volume:carbon conversions from the literature. Eucaryotic organisms contributed about 20–40% of the total organic carbon flux in both the central Ross Sea polynya and in the western polynya, and groups of organisms differed in contribution to the carbon flux at the different sites. Algal carbon flux ranged from 4.5 to 21.1 mg C m⁻² day⁻¹ and consisted primarily of P. antarctica (cell plus mucus) and diatom carbon at all sites. Different diatom species dominated the diatom flux at different sites. Carbon fluxes of small pennate diatoms may have been enhanced by scavenging, by sinking senescent P. antarctica colonies. Heterotrophic carbon flux ranged from 9.2 to 37.6 mg C m⁻² day⁻¹ and was dominated by athecate heterotrophic dinoflagellate carbon in general and by carbon flux of a particular large athecate dinoflagellate at two sites. Fecal pellet carbon flux ranged from 4.6 to 54.5 mg C m⁻² day⁻¹ and was dominated by carbon from ovoid/angular pellets at most sites. Analysis of fecal pellet contents suggested that large protozoans identified by light microscopy contributed to ovoid/angular fecal pellet fluxes. Carbon flux as a percentage of daily primary production was lowest at sites where P. antarctica predominated in the water column and was highest at sites where fecal pellet flux was highest. This indicates the importance of grazers in carbon export.     

Net Zooplankton and the biological pump off Sanriku, Japan

Taxonomic composition, size composition, standing stock, and chemical composition of mesozooplankton were determined to examine the contribution of their fecal pellets to the vertical flux of organic carbon at the outside, the edge, and the center of the warm core ring. The warm core ring significantly affects not only their taxonomic composition and size composition but also their standing stock and chemical composition. The zooplankton at the center of the warm core ring was characterized by the absence of carnivores at the top of the size-trophic relation and filter feeding planktonic tunicates at the bottom. Zooplankton carbon biomass at the outside of the ring was one-third less than that at the center of the ring. The vertical flux of fecal pellets obtained from the pellet volume (12.3 mgC m⁻²d⁻¹) contributed 19 to 96% of the flux (13 to 64 mgC m⁻² d⁻¹) estimated from the body carbon and the fecal pellet production rate. The estimated flux of fecal pellets was 6 to 27% of vertical carbon flux (236 mgC m⁻²d⁻¹) determined by the sediment traps. Microscopic determination of fecal pellets and plankton in the sediment trap samples indicated high grazing activity during the sinking process. Those observations might suggest that particles other than fecal pellets contributed significantly to the vertical carbon flux and fecal pellets were settled directly without loss or being recycled within the surface mixed layer.     

Impacts of the wintertime mesozooplankton community to downward carbon flux in the subarctic and subtropical Pacific Oceans

We compared wintertime depth distributions of the mesozooplankton community and dominant copepods between the subtropical (S1) and subarctic (K2) Pacific Oceans to evaluate the relative importance of actively transported carbon by vertical migrants to sinking particulate organic carbon flux. Primary production was higher and the ratio of sinking particulate organic carbon flux to primary production was lower at S1 compared with those at K2. The mesozooplankton community was lower in abundance and biomass at S1 compared to K2. Copepods were the dominant group among both mesozooplankton abundance and biomass throughout the water column down to 1000 m at both sites. The depth distribution showed that diel vertical migration was obvious for the mesozooplankton abundance and biomass at S1 but was not apparent for the abundance at K2, because the dominant component was diurnally migrating species at S1 and overwintering species residing at mesopelagic depths at K2. The major components of diel migrants were copepods and euphausiids at S1 and only euphausiids at K2. Respiratory flux by the diurnally migrating mesozooplankton was estimated to be 2 mgC m⁻² day⁻¹ at S1 and 7 mgC m⁻² day⁻¹ at K2. The respiratory flux was equivalent to 131% of sedimentary fecal pellet flux at S1 and 136% of that at K2. Because pathways of downward carbon flux are facilitated by the mesozooplankton community, the actively transported carbon (respiration of dissolved inorganic carbon, excretion of dissolved organic carbon and egestion of fecal pellets at depth) might be larger during winter than the flux of sinking fecal pellets.     

Vertical transport of particulate organic matter in the deep Bering Sea and gulf of Alaska

Sediment trap arrays were deployed at two deep ocean stations, one in the Bering Sea and the other in the Gulf of Alaska, in the summer of 1975. The sediment trap was constructed of a pair of polyethylene cylinders (0.185 m² opening) with funnel-shaped bases. The trap is equipped with a lid which is closed before recovery by a tripping messenger system triggered by an electric time release. 37–68% of the total organic carbon fluxes (37–38% in the Bering Sea; 48–68% in the Gulf of Alaska) were represented by large particles (67µm<) such as fecal matter and fecal pellets which contributed minor fractions to the total particulate organic matter concentration in sea water. The total fluxes were 11.1 and 14.2 mg C m^–2d^–1 at 1,510 and 2,610 m respectively at the station (3,800 m) in the Bering Sea, and were 7.60, 4.66 and 3.27 mg C m^–2d^–1 at 900, 1,500 and 1,875 m respectively at the station (4,150 m) in the Gulf of Alaska. The former values are several times greater than the latter, suggesting that there is a regional variation in the vertical carbon flux in deep layers. The fluxes were approximately equivalent to 1 to 3% of primary productivity in the overlying surface layers. These observations suggest that deep-water ecosystems may be influenced by relatively rapid sinking of large particles such as fecal matter and fecal pellets from near surface production.     

Loss Processes of Sinking Fecal Pellets of Zooplankton in the Mesopelagic Layers of the Antarctic Marginal Ice Zone

A time-series sediment trap deployment was carried out in the marginal ice zone (MIZ) of the Antarctic Ocean (64°42′ S, 139°58′E; sea depth of 2930 m), during the austral summer. Cylindrical fecal pellets were the predominant sinking particles at 537 m in the middle of January and most of them disappeared below that depth, the loss of which were 25.3 mg C m⁻² day⁻¹ in the depth range of 537–796 m. Small-sized sinking particles other than fecal pellets increased in that depth range. Analyses of fecal pellets for remnant DNA corresponding to 16S mitochondrial RNA and 28S ribosomal RNA suggested that the large cylindrical fecal pellets at 537 m were produced by Antarctic krill (Euphausia superba) and copepods. According to the presence of the DNA associated with sinking particles, E. superba fecal pellets rapidly disappeared below 537 m, while copepod fecal pellets still remained in the mesopelagic and bathypelagic layers. Small-sized amorphous sinking particles at 537 m also contained E. superba- and copepod-derived DNA. The abundance of trap-collected copepods (Oithona spp. and Oncaea spp.) which are known to be coprophagous increased at 796 m where many fecal pellets disappeared. We suggest that those rapidly sinking pellets were fragmented by copepods with intensified coprorhexy activity (fragmentation of fecal pellets) in the mesopelagic layers, reducing their sinking rates. These smaller and slower sinking particles can be important food sources for detritivorus or coprophagous animals in mesopelagic and bathypelagic layers in the MIZ. This revised version was published online in July 2006 with corrections to the Cover Date.     

Particulate organic carbon supply to the sea bottom: stable carbon isotope ratio analysis of the sediment trap samples at the mouth of otsuchi bay,Northeastern Japan

An array of sediment traps was deployed for the analysis of the pattern of particulate organic carbon (POC) supply to the sea bottom in April, May and July 1988 at the mouth of Otsuchi Bay (about 80 m depth), Northeastern Japan.On the basis of a simple two-component mixing model using stable carbon isotope ratios, the POC flux was separated into marine planktonic and terrestrial components. Both the planktonic and terrestrial POC fluxes had maximum values at 30 m above the sea bottom throughout the three experiments. The planktonic POC flux showed a significant decrease with depth between 30 m and 10 m or 5 m above the bottom. Vertical supply of the planktonic POC and supply of the resuspended planktonic POC were estimated on the basis of regression lines between water depth and the planktonic POC flux in the depth range where the flux decreases with depth.Vertical supply of the planktonic POC and supply of the resuspended planktonic POC to the sea bottom were largest in May (52.1 mgC m^–2 d^–1 and 19.5 mgC m^–2 d^–1 at 5 m above the sea bottom), and horizontal supplies of the terrestrial POC were almost constant (31.9±3.5 mgC m^–2 d^–1 at 5 m above the bottom) throughout the three experiments.     

Contrasts in sedimentation flux below the southern California Current in late 1996 and during the El Niño event of 1997–1998

The vertical flux of particulate matter at 330 m depth in San Lázaro Basin off southern Baja California ranged from 63 to 587 mg m⁻² d⁻¹ between August and November 1996. Organic carbon contents were between 5.6 and 14.8%, yielding flux rates of 9–40 mgC m⁻² d⁻¹. In December 1997 and January 1998, at the height of the strong El Niño event, the respective fluxes (47–202 mg m⁻² d⁻¹ and 3–8 mgC m⁻² d⁻¹) were comparable. The February–June 1998 records, however, revealed sharply reduced mass (1–6 mg m⁻² d⁻¹) and organic carbon (0.2–0.8 mgC m⁻² d⁻¹) fluxes. The organics collected in 1996 were predominantly autochthonous (δ¹³C=−22‰; C/N=8). The variations in δ¹⁵N (8.3–11.0‰) suggest an alternation of new and regenerated production, possibly associated with fluctuations in the intensity of deep mixing during that autumn. The relatively high organic matter fluxes in December 1997 appear to be associated with regenerated production. The average composition from February to June 1998 (δ¹³C=−23.6‰; ¹⁵N=11.7‰; C/N=10.5) indicates degraded material of marine origin. The maximum δ¹⁵N value found (14‰) suggests that deeper, denitrified waters were brought to the surface and possibly advected laterally. Regime changes in the waters of the basin occur at 6–10 week intervals, evidenced by concurrent shifts in most of the measured parameters, including fecal pellet types and metal chemistry. The marine snow-dominated detritus collected showed a shift from a mixed diatom-rich-radiolarian-coccolith assemblage in late 1996 to a coccolith-dominated assemblage, including the contents of fecal pellets, during the 1997–1998 El-Niño period. T–S profiles, plankton analysis and chlorophyll contents of the upper water column indicated that the strong phytoplankton bloom, normally associated with seasonal upwelling along the Pacific coast of Baja, did not occur during the spring of 1998. The persistence of oligotrophic conditions during the 1997–1998 El Niño event favored the dominance of nanoplankton and reduced the vertical flux of particles.     

Sinking Particle Flux in the Subtropical Oligotrophic Northwestern Pacific from a Short-term Sediment Trap Experiment

Time-series sediment traps were deployed in the subtropical oligotrophic northwestern Pacific (SONP) at three depths from August to September 2015 to better understand vertical flux of sinking particles. Sinking particles were collected at 5-day intervals over the sediment trap deployment period. The average total mass flux at water depths of 400 m, 690 m, and 1,710 m was 9.1, 4.4, and 4.1 mg m^-2day^-1, respectively. CaCO₃ materials constituted 50 to 70% of sinking particles while in comparison particulate organic carbon (POC) constituted up to 20%. A synchronous variation of total mass flux was observed at the three depths, indicating that calcite-dominated particles sank from 400 to 1,710 m within a 5-day period. POC flux at these water depths was 2.4, 0.38, and 0.31 mg m^-2day^-1, respectively. Our results indicate low transfer efficiencies of 16% from 400 to 690 m and 13% for the 400 to 1,710 m depth range. The estimated transfer efficiencies were significantly lower than those observed at the K2 station in the northwest Pacific subarctic gyre, presumably because of a prevalence of pico-cyanobacteria in the SONP. Because cyanobacteria have a semi-permeable proteinaceous shell, they are more readily remineralized by bacteria than are siliceous phytoplankton in the northwest Pacific subarctic gyre. Continued surface water warming and expansion of the SONP will likely have a profound impact on ocean acidification in the northwest Pacific, possibly affecting the transfer efficiency of sinking POC to the deep-sea.     

Biogenic debris from the pelagic tunicate, Oikopleura dioica, and its role in the vertical transport of a transuranium element

The accumulation and retention of ²⁴¹Am by the pelagic tunicate Oikopleura dioica were examined using laboratory cultures and radiotracer methodology. Animals (i.e., trunks and tails) and discarded empty houses accumulated Am from seawater, giving volume/volume concentration factors of 59±8 and 10±1, respectively. The half-time for retention of Am in empty labelled houses transferred to non-contaminated seawater was 29 h; the retention half-time of Am in houses discarded by larvaceans feeding on Am-labelled diatoms was 219 h; the half-time of Am in fecal pellets produced by animals feeding on a monospecific diet of diatoms was 134 h, and 247 h for fecal pellets from animals fed a mixed diet. Approximately 30% of filtered cells remained in houses after the houses were discarded. Sinking rates of discarded houses and fecal pellets were found to vary with temperature and size, ranging from 26–157 m day^?1 (house) and from 25–166 m day^?1 (fecal pellets). The ubiquity and abundance of appendicularians, together with their prodigious production of houses (e.g., 10±2 houses day^?1 at 17°C for each experimental animal) point to their potential significance in the vertical transport of Am, and probably other reactive metals, to intermediate depths in the ocean.     

Role of Zooplankton in the Vertical Mass Flux in the Kara and Laptev Seas in Fall

The role of zooplankton in the vertical mass flux in the Kara and Laptev seas was studied during cruise 63 of the R/V Akademik Mstislav Keldysh in August–October 2015. Mass fluxes were estimated using sediment trap samples. The maximum values of the total vertical flux (19600 mg m^?2 day^?1) and particulate organic carbon (POC) flux (464 mg C m^?2 day^?1) were measured close to the Lena River Delta in the Laptev Sea. In the Kara Sea, the total flux (80–2700 mg m^?2 day^?1) and the POC flux (17–130 mg C m^?2 day^?1) were substantially higher than the estimates published earlier. The fecal pellet flux varied from 2 to 92 mg C m^?2 day^?1 and made up 4–190% of the total organic carbon flux. The mineral composition of fecal pellets largely mirrored that of suspended particulate matter. Clay minerals in the fecal pellets were more abundant than in particulate matter in the areas with noticeable freshwater impact. The flux of zooplankton carcasses varied from 0.1 to 66.4 mg C m^?2 day^?1 and made up 0.2–72% of total POC flux. The results are discussed in relation to the abundance and composition of zooplankton, the concentration and composition of suspended particulate matter, hydrophysical conditions, and methods of sample preparation for analysis.     

Role of large particles in the transport of elements and organic compounds through the oceanic water column

During the past decade data from a variety of sources have been obtained which show conclusively that the relatively rare, large particles sinking through the water column are responsible for the majority of the downward vertical mass flux in the sea. This finding has important implications for understanding the transfer, distribution and fate of elements and organic compounds in marine waters. The “large” (> 100 μm) detrital particles responsible for vertical flux are primarily biogenic and range in size and composition from small, discrete fecal pellets and plankton hard parts to large, amorphous aggregates or “snow” which contain both organic and inorganic constituents. Depending on size, shape and density, these particles sink at rates ranging from <1 to >1000 m day⁻¹. Several methods have been developed for sampling these particles of which in situ sediment trapping has probably furnished the most comprehensive qualitative and quantitative information on the role large particles play in material transport. Flux studies have highlighted the importance of marine heterotrophs in packaging fine, suspended particulate matter into large rapidly sinking particles which accelerate the movement of incorporated materials to depth. Large particle production via biological packaging is not restricted to the euphotic zone but can occur at all depths and information is now accruing on rates of production of large particles in the water column. Chemical analyses of sedimenting particles collected in sediment traps and those sampled by other means have allowed quantifying vertical fluxes and residence times of elements, radionuclides and organic compounds (natural and anthropogenic) in various oceanic regimes. Pertinent studies dealing with the above aspects are reviewed and several areas for future research are suggested.     

Distributions and upward fluxes of particulate matter near bottom in the southeastern Bering Sea Shelf

Time-series measurements of temperature, salinity, suspended matter and beam attenuation coefficient () were measured at four hour intervals for about two days in June/ July 1982 in the middle shelf region and the coastal region of the southeastern Bering Sea. Current meters were also moored at the same locations.Depth-time distributions of indicated that profiles of suspended matter resulted from a combined process of resuspension of underlying sediments and sinking of suspended particles. Average-values for all measurements for particles revealed that the upward transport of particles due to resuspension formed a boundary layer, with a thickness apparently related to scalar speed. The average-profiles of the particle volume concentration were assumed to result from a balance between the sinking and diffusive flux of particles under a steady state, and the upward fluxes were calculated. Within the boundary layer, values of the upward fluxes of particulate organic matter linearly decreased with the logarithm of distance from the bottom. Fluxes of organic carbon at the upper edge of the boundary layer were 0.375 gC·m^–2·day^–1 in the middle shelf region (18 m above the bottom, bottom depth=78m) and 0.484gC·m^–2·day^–1 in the coastal region (25 m above the bottom, bottom depth=33m), and fluxes of nitrogen in both regions were 0.067 gN·m^–2·day^–1. The flux of organic carbon obtained in the middle shelf region (18 m above the bottom) agreed approximately with the flux (0.416 gC·m^–2·day^–1) calculated by substituting primary production data into the empirical equation of Suess (1980).     

Seasonal changes and biochemical composition of the labile organic matter flux in the Cretan Sea

Downward fluxes of labile organic matter (lipids, proteins and carbohydrates) at 200 (trap A) and 1515 m depth (trap B), measured during a 12 months sediment trap experiment, are presented, together with estimates of the bacterial and cyanobacterial biomasses associated to the particles. The biochemical composition of the settling particles was determined in order to provide qualitative and quantitative information on the flux of readily available organic carbon supplying the deep-sea benthic communities of the Cretan Sea. Total mass flux and labile carbon fluxes were characterised by a clear seasonality. Higher labile organic fluxes were reported in trap B, indicating the presence of resuspended particles coming from lateral inputs. Particulate carbohydrates were the major component of the flux of labile compounds (on annual average about 66% of the total labile organic flux) followed by lipids (20%) and proteins (13%). The biopolymeric carbon flux was very low (on annual average 0.9 and 1.2 gC m⁻² y⁻¹, at trap A and B). Labile carbon accounted for most of the OC flux (on annual average 84% and 74% in trap A and B respectively). In trap A, highest carbohydrate and protein fluxes in April and September, corresponded to high faecal pellet fluxes. The qualitative composition of the organic fluxes indicated a strong protein depletion in trap B and a decrease of the bioavailability of the settling particles as a result of a higher degree of dilution with inorganic material. Quantity and quality of the food supply to the benthos displayed different temporal patterns. Bacterial biomass in the sediment traps (on average 122 and 229 μgC m⁻² d⁻¹ in trap A and B, respectively) was significantly correlated to the flux of labile organic carbon, and particularly to the protein and carbohydrate fluxes. Cyanobacterial flux (on average, 1.1 and 0.4 μgC m⁻² d⁻¹, in trap A and B, respectively) was significantly correlated with total mass and protein fluxes only in trap A. Bacterial carbon flux, equivalent to 84.2 and 156 mgC m⁻² y⁻¹, accounted for 5–6.5% of the labile carbon flux (in trap A and B respectively) and for 22–41% protein pool of the settling particles. These results suggest that in the Cretan Sea, bacteria attached to the settling particles represent a potential food source of primary importance for deep-sea benthic communities.     

Functional roles of interzonal migrating mesozooplankton in the western subarctic Pacific

Grazing experiments and production estimation based on life-history analysis of Neocalanus copepods (N. cristatus, N. plumchrus and N. flemingeri) were carried out in the Oyashio region to understand the carbon flows associated with the interzonal migrating copepods. These copepods, and also Eucalanus bungii, fed on nano- and micro-sized organisms non-selectively throughout the season. However, diatoms were the dominant food resource until May and organisms, such as ciliates were the major resource after May. Daily growth rate was estimated from the Ikeda–Motoda, Huntley–Lopez and Hirst–Sheader models. Since the growth rates were considered to be overestimates for the Huntley–Lopez model and underestimates for the other two models, we applied the weight-specific growth rates previously reported for these species in the Bering Shelf. Surface biomass of Neocalanus increased rapidly in June during the appearance of C5, and a successive increase of overwintering stock was evident in the deeper layer. The deep biomass decreased gradually from September to May during the dormant and reproduction period. N. cristatus has the largest annual mean biomass (2.3 gC m⁻²), followed by N. plumchrus (1.1) and N. flemingeri (0.4). Daily production rate of Neocalanus varied from 0.4 to 363.4 mgC m⁻² day⁻¹, to which N. cristatus was the largest contributor. Annual production was estimated as 11.5 gC m⁻² year⁻¹ for N. cristatus, 5.7 for N. plumchrus and 2.1 for N. flemingeri, yielding annual P/B ratio of 5 for each species. The annual production of Neocalanus accounted for 13.2% of the primary production in the Oyashio region. Their fecal pellets were estimated to account for 14.9% (0.7 gC m⁻² year⁻¹) of sinking flux of organic carbon at 1000-m depth. Moreover, their export flux by ontogenetic vertical migration, which is not measured by sediment trap observations, is estimated to be 91.5% (4.3 gC m⁻² year⁻¹) of carbon flux of sinking particles at 1000-m depth. These results suggest the important role of interzonal migrating copepods in the export flux of carbon.     

Vertical flux of biogenic matter during a Lagrangian study off the NW Spanish continental margin

Lagrangian experiments with short-term, drifting sediment traps were conducted during a cruise on RRS Charles Darwin to the NW coast of Spain to study the vertical flux and composition of settling biogenic matter. The cruise was split into two legs corresponding to (i) a period of increased production following an upwelling event on the continental shelf (3–10 August 1998) and (ii) an evolution of a cold water filament originating from the upwelled water off the shelf (14–19 August). The export of particulate organic carbon (POC) from the upper layer (0–60m) on the shelf was 90–240mgC.m⁻².d⁻¹ and off the shelf was 60–180mgC.m⁻².d⁻¹. Off shelf the POC flux at 200m was 50–60mg.m⁻².d⁻¹. A modest sedimentation of diatoms (15–30mgC.m⁻².d⁻¹) after the upwelling was associated with increased vertical flux of chlorophyll a (1.8–2.1mg.m⁻².d⁻¹) and a decrease of the POC:PON molar ratio of the settled material from 9 to 6.4. Most of the pico-, nano-, and microplankton in the settled material were flagellates; diatoms were significant during the on shelf and dinoflagellates during the off shelf leg. Off shelf, the exponential attenuation of POC flux indicated a strong retention capacity of the plankton community between 40 and 75m. POC:PON ratio of the settled particulate matter decreased with depth and the relative portion of flagellates increased, suggesting a novel, flagellate and aggregate mediated particulate flux in these waters. Export of POC from the euphotic layer comprised 14–26% of the integrated primary production per day during the on shelf leg and 25–42% during the off shelf leg, which characterises the importance of sedimentation in the organic carbon budget of these waters.     

High-frequency fluxes of labile compounds in the central Ligurian Sea,northwestern Mediterranean

Sinking particles were collected every 4 h with drifting sediment traps deployed at 200 m depth in May 1995 in a 1-D vertical system during the DYNAPROC observations in the northwestern Mediterranean sea. POC, proteins, glucosamine and lipid classes were used as indicators of the intensity and quality of the particle flux. The roles of day/night cycle and wind on the particle flux were examined. The transient regime of production from late spring bloom to pre-oligotrophy determined the flux intensity and quality. POC fluxes decreased from, on average, 34 to 11 mg m⁻² d⁻¹, representing 6–14% of the primary production under late spring bloom conditions to 1–2% under pre-oligotrophic conditions. Total protein and chloroplast lipid fluxes correlated with POC and reflected the input of algal biomass into the traps. As the season proceeded, changes in the biochemical composition of the exported material were observed. The C/N ratio rose from 7.8 to 12. Increases of serine (10–28% of total proteins), total lipids (7–9 to 14–28% of POC) and reserve lipids (1–5 to 5–22% of total lipids) were noticeable, whereas total protein content in POC decreased (20–27 to 18–7%). N-acetyl glucosamine, a tracer of fecal pellet flux, showed that zooplankton grazing was a major vector of downward export during the decaying bloom. Against this background pattern, episodic events specifically increased the flux, modifying the quality and the settling velocity of particles. Day/night signals in biotracers (POC, N-acetyl glucosamine, protein and chloroplast lipids) showed that zooplankton migrations were responsible for sedimentation of fresh material through fast sinking particles (V=170–180 m d⁻¹) at night. Periodic signatures of re-processed material (high lipolysis and bacterial biomass indices) suggested that other zooplankton fecal pellets or small aggregates, probably of lower settling velocities (V<170 m d⁻¹), contributed to the flux during calm periods. At the beginning of the experiment, during the development of a prymnesiophyte bloom in the upper layers, the sterol signal with no periodicity enabled us to estimate high particle settling velocities (⩾600 m d⁻¹) likely related to large aggregate formation. A wind event increased biotracer fluxes (POC, protein, chloroplast lipids). The rapid transmission of surface signals through extremely fast sinking particles could be a general feature of particle fluxes in marine areas unaffected by horizontal advection.     

Effects of the copepod community structure on fecal pellet flux in Kagoshima bay,a deep,semi-enclosed embayment

Seasonal changes in the shape and size composition of fecal pellets were investigated with sediment trap samples from 50 and 150 m in Kagoshima Bay to evaluate how the mesozooplankton community affects fecal pellet flux. Deep vertical mixing was evident in March, and thermal stratification was developed above 50 m in June, August and November. Chlorophyll a, suspended particulate organic carbon (POC) and copepod abundance were uniform throughout the water column during the seasonal mixing and concentrated above 50 m in the stratified seasons. Calanoids were the most predominant copepods in March and poecilostomatoids composed more than 45% of the copepod community in June, August and November. Fecal pellet fluxes at 50 and 150 m were the highest in March, nearly half of POC flux. The relative contribution declined considerably in the other months, especially for less than 4% of POC flux in August. The decline was corresponded to the predominance of cyclopoids and poecilostomatoids. Cylindrical pellets dominated the fecal matters at both depths throughout the study period, while larger cylindrical pellets nearly disappeared at 150 m in June, August and November. Copepod incubation revealed that cylindrical and oval pellets were egested by calanoids and the other copepods, respectively. We suggest that cylindrical fecal pellets produced by calanoid copepods contribute to feces flux but the predominance of poecilostomatoids and/or cyclopoids decreases feces flux via the increase of oval pellets and fragmentation of larger cylindrical pellets.     

Export flux of particles at the equator in the western and central Pacific ocean

Export of particles was studied at the equator during an El Nin˜o warm event (October 1994) as part of the French ORSTOM/FLUPAC program. Particulate mass, carbon (organic and inorganic) (C), nitrogen (N), and phosphorus (P) export fluxes were measured at the equator in the western and central Pacific during two 6–7 day-long time-series stations located in the warm pool (TS-I at 0°, 167°E) and in the equatorial HNLC situation (TS-II at 0°, 150°W), using drifting sediment traps deployed for 48 h at four depths (between, approximately, 100 and 300 m).The particulate organic carbon (POC) fluxes at the base of the euphotic zone (0.1 % light level), were approximately four times lower at TS-I than at TS-11 (4.1 vs. 17.0 mmol C m^-2 day^-1). Conversely, fluxes measured at 300 m were similar at both sites (3.6vs. 3.7 mmol C m⁻² day⁻¹ at TS-I and TS-11, respectively). This change in export fluxes was in good agreement with food-web dynamics in the euphotic zone characterized by an increase in plankton biomasses and metabolic rates and a shift towards larger size from TS-1 to TS-II. The POC flux profiles indicated high remineralization (up to 78%) of the exported particles at TS-II, between 100 and 200 m in the Equatorial Undercurrent. According to zooplankton ingestion estimates from 100 – 300 m, 60% of this POC loss could be accounted for by zooplankton grazing. At TS-I, no marked increase of flux with depth was observed, and we assume that loss of particles was compensated by in-situ particle production by zooplankton. Fluxes of particulate nitrogen and phosphorus followed the same general patterns as the POC fluxes. The elemental and pigment composition of the exported particles was not very different between the two stations. In particular, the POCYN flux molar ratio at the base of the euphotic zone was low, 6.9 and 6.2 at TS-1 and TS-II, respectively.For particulate inorganic carbon (mainly carbonate) flux, values at the base of the euphotic zone averaged 0.9 mmol C m-2 day-1 at TS-I and 2.3 mmol C m-2 day-1 at TS-11 (corresponding to a 2.6-fold increase) and showed low depth changes at both stations.POC export flux (including active flux associated with the interzonal migrants) at the 0.1 % light level depth represented only 8% of primary production (1°C uptake) measured at TS-1 and 19% at TS-II. For the time and space scales considered in the present study, new primary production, as measured by the 15N method, was in good agreement with the total export flux in the HNLC situation, thus leading to negligible dissolved organic carbon (DOC) or nitrogen (DON) losses from the photic zone. Conversely, export flux was found to be only 50% (C units) and 60% (N) of new production in the oligotrophic system, either because of an overestimation by the 15N method or of a significant export of DOC and DON.Comparison with other oceanic regions shows that export flux in the warm pool was within the same range as in the central gyres. On the other hand, comparison with EgPac data in the central Pacific suggests that there is no straightforward relation between the magnitude of the export and surface nitrate concentrations.     

Biogeochemistry of organic matter in the Laurentian Trough, I. Composition and vertical fluxes of rapidly settling particles

Settling particles from duplicate free-drifting sediment traps were collected at 150 m depth in May and July at a landward and a seaward site in the 350 m deep Laurentian Trough. The total organic carbon (TOC) fluxes were high (95–454 mg/m²/d), comparable to those reported for Dabob Bay (a similar moderately productive deep coastal environment) and for the highly productive Peru upwelling region. The TOC (26–67 mg C/g) consisted of lipids (17–37%), carbohydrates (7.9–16%), hydrolysable amino acids (8.4–16%), labile proteins (0.3–2.6%), and a non-characterized fraction (40–64%). Amino acids, proteins and uncharacterized compounds accounted for 24–42, 1–10 and 58–76%, respectively, of total nitrogen (2.3–7.7 mg N/g). The pigment fraction was largely dominated by pheopigments (0.06-1.15 mg/g vs 0.004–0.15 mg/g for chlorophyll a). C/N and C/pigment ratios indicated that on average, about half of the carbon flux was of terrigenous origin. Marine sources included a dominant zooplanktonic contribution, indicated by the abundance of fecal pellets, lipids and pheopigments, and a smaller contribution from fresh algae. Cluster and correlation analyses confirmed the decoupling of pigment and TOC fluxes and the strong zooplanktonic influence of the trap material. Despite large day-to-day and inter-trap variability, clear differences were observed in the fluxes, TOC content and composition at both sampling sites and months. Such trends are attributed to the relative contribution from terrestrial and marine sources and seasonal patterns of primary production.     

Downward fluxes of settling particles in the deep Cretan Sea (NE Mediterranean)

During the CINCS project (Pelagic–benthic Coupling IN the oligotrophic Cretan Sea—NE Mediterranean), a single mooring with two sediment traps (at 200 and 1515m water depth) and two current meters was deployed in the southern Cretan Sea margin at a depth of 1550 m. A second mooring deployed at the 500 m station was lost, as a result of fishing activities. The duration of the study was 12 months (November 1994 to November 1995) with sampling intervals of 15 or 16 days. The traps were retrieved, serviced and the sedimented material was collected every 6 months. In total, 48 samples were collected (24 from each trap) throughout the study period and fluxes of total particulate mass, opal, organic matter, carbonates, and lithogenic component were measured. Natural radionuclides (²¹⁰Po and ²¹⁰Pb) were determined for all trap samples. Total mass flux and the fluxes of four major constituents increased with depth, the total mass flux reaching values of nearly 550 mg m⁻² d⁻¹ at 1515 m and 187 mg m⁻² d⁻¹ at 200 m depth, following the same patterns observed in other experiments (ECOMARGE, SEEP-I, SEEP-II). The mean annual mass fluxes were 209 and 49.8 mg m⁻² d⁻¹ at the near bottom and near surface trap respectively. This suggests that lateral transport of particulate matter is of importance in the area. Total mass fluxes at the two depths were characterized by different seasonal fluctuations, although a general decreasing trend was observed from the I (winter) to the II (summer) deployment at both depths. This was mainly a result of reductions in aluminosilicate inputs during the summer dry period. At 200 m depth carbonates were more important during winter, because of a large carbonate input consisting mainly of coccoliths of Emiliania huxleyi, while during the summer decreased fluxes of carbonates and aluminosilicates resulted in a reduction of the mass flux. In contrast, at 1515 m depth the lithogenic component was the dominant component during the winter deployment, indicating a terrigenous input. During the summer period the decrease in mass flux was strongly effected by the decrease in aluminosilicates. There was a diminution in the organic carbon content with a concomitant increase in total mass flux, which, together with the almost negligible increase in the annual ²¹⁰Pb activity with depth and the increase of ²¹⁰Po activity with depth could be interpreted as indicating a contribution of resuspended material to the input at 1515 m. The complex mesoscale circulation of the Cretan Sea, consisting of a cyclone (east)–anticyclone (west) system, controls particle transfer in the area. This hydrodynamic system seems to move water masses towards the southern Cretan Sea margin, and consequently carry materials from the open sea to the upper slope and shelf.