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.     

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.     

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.     

Seasonal changes in the mesozooplankton biomass and community structure in subarctic and subtropical time-series stations in the western North Pacific

Seasonal changes in mesozooplankton biomass and their community structures were observed at time-series stations K2 (subarctic) and S1 (subtropical) in the western North Pacific Ocean. At K2, the maximum biomass was observed during the spring when primary productivity was still low. The annual mean biomasses in the euphotic and 200- to 1000-m layers were 1.39 (day) and 2.49 (night) g C m^?2 and 4.00 (day) and 3.63 (night) g C m^?2, respectively. Mesozooplankton vertical distribution was bimodal and mesopelagic peak was observed in a 200- to 300-m layer; it mainly comprised dormant copepods. Copepods predominated in most sampling layers, but euphausiids were dominant at the surface during the night. At S1, the maximum biomass was observed during the spring and the peak timing of biomass followed those of chlorophyll a and primary productivity. The annual mean biomasses in the euphotic and 200- to 1000-m layers were 0.10 (day) and 0.21 (night) g C m^?2 and 0.47 (day) and 0.26 (night) g C m^?2, respectively. Copepods were dominant in most sampling layers, but their mean proportion was lower than that in K2. Mesozooplankton community characteristics at both sites were compared with those at other time-series stations in the North Pacific and with each other. The annual mean primary productivities and sinking POC fluxes were equivalent at both sites; however, mesozooplankton biomasses were higher at K2 than at S1. The difference of biomasses was probably caused by differences of individual carbon losses, population turnover rates, and trophic structures of communities between the two sites.     

The downward flux of biogenic material in the NE subarctic Pacific: importance of algal sinking and mesozooplankton herbivory

In the present study we examine factors that affect the downward flux of biogenic carbon in the NE subarctic Pacific, one of the important high-nutrient-low-chlorophyll (HNLC) regions in the open ocean. We focus on the role of mesozooplankton, since their seasonal peaks in biomass and growth are in phase with the seasonal variations in the downward POC fluxes, whereas phytoplankton biomass is more-or-less uniform year-round. The relative importance of mesozooplankton and algal sinking was examined using the pigment composition of material accumulated in short-term free-drifting sediment traps positioned just below the upper stratified surface layer (ca. 100–200 m). This was compared with the phytoplankton composition in the surface waters, and with the grazing activity (gut pigments and fecal pellet production rates) of the most abundant large copepods. We also examined whether the relationships between the downward flux of carbon and pelagic processes were similar in the coastal, continental margin and offshore HNLC regions of the NE subarctic Pacific, the latter represented by Ocean Station Papa (OSP).Our results show that grazing had a variable impact on the downward flux of biogenic carbon. Carbon-transformed pheopigments (particularly pyropheophorbide a, frequently associated with copepod grazing) represented up to 13% of the total downward POC flux inshore (in May 1996) and 8–9% at OSP in May and February 1996, respectively. This flux of pheopigments was accompanied by a large potential input of fecal pellets from large copepods (as estimated from defecation rates of freshly collected animals) only in May 1996 at OSP, suggesting that pheopigments came from other sources (other herbivores, senescing algae) in February. The larger flux of pheopigments in May was probably related to the abundance of mesozooplankton at that time of the year. During summer (August 1996), both the flux of pheopigments and the potential input of fecal pellets from large copepods were negligible at OSP, consistent with more intense pelagic recycling reported in other studies. Inshore, the flux of carbon-transformed pheopigments was slightly higher than at OSP, and its contribution to the downward POC flux in May 1996 was twice that in August 1996. In contrast, the potential input of feces carbon was higher in August than in May 1996, again suggesting other sources for pheopigments found in the traps. The contribution of sinking phytoplankton to the downward biogenic flux was negligible in summer, when prymnesiophytes (indicated by the presence of 19′-hexanoyloxyfucoxanthin) and pelagophytes (19′-butanoyloxyfucoxanthin-containing) dominated in surface offshore waters. The contribution of sinking algae was maximal (9%) in winter (February 1996) at OSP, when fucoxanthin (mainly a diatom marker) dominated the carotenoid composition in the traps and when the abundance of diatoms in surface waters showed its seasonal maximum for this station. Inshore, the low contribution of diatoms (fucoxanthin) to the sinking fluxes may have resulted from inadequate sampling (i.e. the spring bloom may have been missed).Overall, we conclude that: (a) large copepods significantly influenced the downward POC flux only during spring at OSP; (b) unidentified herbivores (e.g. salps, pteropods) producing pigmented, fast-sinking fecal material likely had an important impact during winter; (c) algal sinking made a small contribution to the downward POC flux (maximum in winter); and (d) neither algal sinking nor mesozooplankton grazing had a significant influence on the downward flux of biogenic material in summer at OSP.     

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.     

北太平洋亚热带环流区与西太平洋暖池区浮游动物群落垂直分布的差异及其对碳通量的意义

The mesozooplankton in both epipelagic and mesopelagic zones is essentially important for the study of ecosystem and biological carbon pump. Previous studies showed that the diel vertical migration(DVM) pattern of mesozooplankton varied among ecosystems. However, that pattern was largely unknown in the Western Pacific Warm Pool(WPWP). The vertical distribution, DVM and community structure of mesozooplankton from the surface to 1 000 m were compared at Stas JL7K(WPWP) and MA(North Pacific Subtropical Gyre, NPSG). Two sites showed similarly low biomass in both epipelagic and mesopelagic zones, which were in accordance with oligotrophic conditions of these two ecosystems. Stronger DVM(night/day ratio) was found at JL7K(1.31) than that at MA(1.09) on surface 0–100 m, and an obvious night increase of mesopelagic biomass was observed at JL7K, which was probably due to migrators from bathypelagic zone. Active carbon flux by DVM of zooplankton was estimated to be 0.23 mmol/(m2·d) at JL7K and 0.16 mmol/(m~2·d) at MA. The community structure analysis showed that calanoid copepods, cnidarians and appendicularians were the main contributors to DVM of mesozooplankton at both sites. We also compared the present result with previous studies of the two ecosystems,and suggested that the DVM of mesozooplankton was more homogeneous within the WPWP and more variable within the NPSG, though both ecosystems showed typically extremely oligotrophic conditions. The different diel vertical migration strength of mesozooplankton between NPSG and WPWP implied different efficiency of carbon pump in these two ecosystems.     

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.     

Downward flux of zooplankton fecal pellets in the upper 2,000 m water column of the Izu Trench,western North Pacific

Sinking particulate matter were obtained from twelve depths using free-drifting sediment trap arrays which were deployed in the upper 2,000 m water column of the Izu Trench, northwest Pacific Ocean. The largest flux of 146 mgC m^–2 day^–1 was observed at 150 m depth. The flux generally decreased with depth below the maximum, however, minor flux peaks occurred at 1,000 and 1,250 m depth (>30 mgC m^–2 day^–1). Sinking large particles (>100 µm) were composed of fecal pellets typical of crustaceans, macroscopic aggregates, and planktonic organisms and their fragments. Three major components constituted 19%, 20% and 29%, respectively, of the total carbon flux (averaged from the fluxes at five depths; 50, 100, 150, 1,000 and 2,000 m). Among them, fecal pellet flux and large organism flux were well correlated with the total flux. The close correspondence between the fecal flux and the total carbon flux suggests that the latter is derived from a group of variables including other biogenic matter, among which fecal pellet is one of the leading factors controlling total flux, though the latter is only a minor covariable in quantity. Vertical flux profiles of fecal pellets and their internal constituents revealed some new inputs of feces occurring through the water column. This phenomenon implies that downward transportation of organic material is characterized by feeding and egestion activities of zooplankton, including overlapping processes of sinking and dispersion of large fecal particles and repackaging of dispersed small particles.     

The distribution and vertical flux of fecal pellets from large zooplankton in Monterey bay and coastal California

We sampled zooplankton and fecal pellets in the upper 200 m of Monterey Bay and nearby coastal regions in California, USA. On several occasions, we observed high concentrations of large pellets that appeared to be produced during night-time by dielly migrating euphausiids. High concentrations of pellets were found in near-surface waters only when euphausiids co-occurred with high concentrations of large (>10 μm) phytoplankton. Peak concentrations of pellets at mid-depth (100 or 150 m) during the day were consistent with the calculated sinking speeds of pellets produced near the surface at night. At these high flux locations (HI group), pellet concentrations declined below mid-depth. In contrast, at locations where the phytoplankton assemblage was dominated by small phytoplankton cells (<10 μm), pellet production and flux were low (LO group) whether or not euphausiid populations were high. Protozooplankton concentrations did not affect this pattern. We concluded that the day and night differences in pellet concentration and flux in the HI profiles were mostly due to sinking of dielly-pulsed inputs in the surface layer, and that small zooplankton (Oithona, Oncaea), heterotrophic dinoflagellates, and bacterial activity probably caused some pellet degradation or consumption below 100 m. We estimated that consumption of sinking pellets by large copepods was insignificant. High fluxes of pellets were episodic because they required both high concentrations of large phytoplankton and large stocks of euphausiids. Under these conditions, flux events overwhelmed retention mechanisms, resulting in large exports of organic matter from the upper 200 m.     

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.     

Elevated biomass of mesozooplankton and enhanced fecal pellet flux in cyclonic and mode-water eddies in the Sargasso Sea

Accumulating evidence points to the importance of mesoscale eddies in supplying nutrients to surface waters in oligotrophic gyres. However, the nature of the biological response and its evolution over time has yet to be elucidated. Changes in mesozooplankton community composition due to eddy perturbation also could affect biogeochemical cycling. Over the course of two summers we sampled seven eddies in the Sargasso Sea. We focused on and followed a post-phytoplankton bloom cyclonic eddy (C1) in 2004 and a blooming mode-water anticyclonic eddy (A4) in 2005. We collected zooplankton in all eddies using a Multiple Opening and Closing Net Environmental Sampling System (MOCNESS) and quantified biomass (>0.15 mm, in five size fractions) from 0 to 700 m over nine discrete depth intervals. Zooplankton biomass (>0.5 mm) in the upper 150 m was similarly enhanced at night for the periphery of C1 and the center of A4 at 0.514 g m⁻² and 0.533 g m⁻², respectively, compared to outside (0.183 g m⁻² outside C1 and 0.197 g m⁻² outside A4). Despite minimal chlorophyll a enhancement and dominance by picoplankton in C1, zooplankton biomass increased most for the largest size class (>5 mm). Gut fluorescence for euphausiids and large copepods was also elevated on the C1 periphery. In A4, peak biomass occurred at eddy center coincident with peak primary production, but was highly variable (changing by >3-fold) over time, perhaps resulting from the dense, but patchy distribution of diatom chains in this region. Shifts in zooplankton community composition and abundance were reflected in enhancement of fecal pellet production and active transport by diel vertical migration in eddies. Inside C1 the flux of zooplankton fecal pellets at 150 m in June 2004 was 1.5-fold higher than outside the eddy, accounting for 9% of total particulate organic carbon (POC) flux. The flux of fecal pellets (mostly from copepods) increased through the summer in eddy A4, matching concurrent increases in zooplankton <2 mm in length, and accounting for up to 12% of total POC flux. Active carbon transport by vertically migrating zooplankton was 37% higher on the periphery of C1 and 74% higher at the center of A4 compared to the summer mean at the Bermuda Atlantic Time-series Study (BATS) station. Despite contrasting responses by the phytoplankton community to cyclonic and mode-water eddies, mesozooplankton biomass was similarly enhanced, possibly due to differential physical and biological aggregation mechanisms, and resulted in important zooplankton-mediated changes in mesoscale biogeochemistry.     

Feeding,egg production,and respiration rate of pteropods <Emphasis Type="Italic">Limacina</Emphasis> in Arctic seas

The feeding, egg production, and respiration rate of the dominant pteropod Limacina helicina have been studied in Russia’s Arctic seas. The sinking rates of fecal pellets and dead individuals have been measured to estimate their role in vertical carbon flux. As has been shown, the rate of ecophysiological processes taking place in the pteropods is higher than that of copepods, the main consumers of phytoplankton. The gut pigment content in Limacina (3084 ng ind^–1 as a maximum) was two orders of magnitude higher than in copepods. The egg production rate in Limacina even without feeding reached 4000 eggs ind^–1 versus 350–450 egg ind^–1 typical of the dominant copepods even with excess food. A close correlation between the pteropod feeding rate and individual body weight was observed for Limacina rather than a correlation with food concentration. The experimentally estimated sinking rate of Limacina fecal pellets was 270 m day^–1, higher than for most copepods. The sinking rate of dead pteropods reaches 2000 m day^–1. According to the literature, discarded mucous feeding nets sink at a rate of 80 to 1080 m day^–1. Evidently, pteropods play a significant role in biogeochemical cycles by accelerating sedimentation. High rates of all studied processes suggest that Limacina are an important component of plankton communities and play the most important role in trophodynamics at sites of their accumulation.     

Interannual variability in vertical export in the Ross Sea: Magnitude, composition, and environmental correlates

The vertical flux of particulate matter from the surface of the Ross Sea, Antarctica, has been suggested as being large, with substantial seasonal and spatial variations. We conducted a study in which vertical flux was quantified using sediment traps deployed at 200 m and compared to estimates calculated from one-dimensional budgets of nutrients (nitrogen and silicon). Estimates of flux were collected at two locations in the southern Ross Sea from late December to early February during four years: 2001-2002, 2003-2004, 2004-2005, and 2005-2006. Phytoplankton biomass and vertical flux varied substantially seasonally and spatially between the two sites, and among years. The greatest flux was observed in 2001-2002, with a short-term maximum organic carbon flux of 3.13 mmol m⁻² d⁻¹, and the summer mean organic carbon flux equal to 0.93 mmol m⁻² d⁻¹. In contrast, the mean carbon flux at the same site in 2003-2004 was over an order of magnitude less, averaging 0.19 mmol m⁻² d⁻¹, despite the fact that productivity in that year was substantially greater. In 2005-206 the contribution of fecal pellets to flux was smallest among all years, and the pellet contribution ranged from <1 to more than 50% of organic flux. As the moorings also had surface layer fluorometers, the relationship between surface biomass and sediment trap flux was compared. Temporal lags between surface fluorescence and flux at 200 m maxima in 2003-2004 and 2004-2005 ranged from two to six days; however, in 2005-2006 the temporal offset between biomass and flux was much longer, ranging from 11 to 27 days, suggesting that fecal pellet production appeared to increase the coupling between flux and surface production. Estimates of export from the upper 200 m based on one-dimensional nutrient budgets were greater than those recorded by the sediment traps. Nutrient budgets also indicated that siliceous production averaged ca. 40% of the total annual production. The variations observed in the flux of biogenic matter to depth in the Ross Sea are large, appear to reflect different forcing among years, and at present are not adequately understood. However, such variability needs to be both understood and represented in biogeochemical models to accurately assess and predict the effects of climate change on biogeochemical cycles.     

Comparison of sinking particles in the upper 200 m between subarctic station K2 and subtropical station S1 based on drifting sediment trap experiments

Drifting sediment trap experiments were conducted during various seasons to elucidate the characteristics of particles sinking through the upper 200 m of the water column in the western Pacific at subarctic station K2 and subtropical station S1. The sinking particle flux increased when primary productivity was high at each station, during June–July at K2 and during February at S1. Biogenic opal (Opal) and CaCO₃ were the major components of the fluxes at K2 and S1, respectively. Contrary to the expectation of a high flux at the eutrophic station K2 and low flux at the oligotrophic station S1, the annual average organic carbon fluxes at 100 m were comparable at both stations: 62.7 mg C m^?2 day^?1 at K2 and 56.1 mg C m^?2 day^?1 at S1. The similarity of the fluxes was perhaps a reflection of the unexpectedly high primary production at S1. At K2, the organic carbon export ratio (organic carbon flux/primary productivity) was significantly and negatively correlated with primary production and tended to decrease with depth. The magnitude of the rate of attenuation of the organic carbon flux with depth was larger at S1 than at K2. This rate of attenuation tended to decrease and increase with primary production at K2 and S1, respectively. The explanation for these patterns may be that the flux of labile organic carbon at relatively shallow depths decreased with increasing primary production at K2, and zooplankton grazing pressure increased with increasing primary productivity at S1.     

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.     

Downward export of carbon by diel migrant mesozooplankton in the central equatorial Pacific

Using data collected during cruises of the JGOFS equatorial Pacific Study in March/April and October of 1992 at the equator (140°W), we examine the downward transport of carbon by three size classes of die] migrant mesozooplankton (200–500 gm, 500–1000 μm and 1000–2000 gm). In addition to respiratory carbon flux, we consider the flux due to mortality of migrators below the euphotic zone. Diel migrant mesozooplankton biomass was estimated from the difference between nighttime and daytime biomass within the euphotic zone. Except for a four-day period early in the March/April cruise, mesozooplankton nighttime biomass was significantly larger than daytime biomass within the euphotic zone during both cruises. We estimate that the downward flux of carbon from the euphotic zone due to mesozooplankton die] vertical migrators was an average of 0.6 mmol Cm⁻² d⁻¹ and 1.1 mmol C m⁻² d⁻¹ during the March/April and October cruises, respectively. Addition of this flux to the gravitational particle sinking flux estimated from²³⁴Th measurements during the same period results in a 31 % increase in the carbon export flux from the euphotic zone in the equatorial Pacific during the March/April cruise and a 44% increase in the October cruise. The migratory flux is strongly dependent on whether feeding takes place below the euphoric zone, the length of time migrators spend in the deep waters, and the mortality rate of migrators.     

Vertical distribution of prokaryote production and abundance in the mesopelagic and bathypelagic layers of the Canada Basin,western Arctic: Implications for the mode and extent of organic carbon delivery

The vertical distributions of prokaryote heterotrophic production (³H-leucine incorporation rate) and abundance were investigated in the meso- and bathy-pelagic layers of the Canada Basin, western Arctic Ocean, during September 2009. Prokaryote production and abundance were high in the Pacific-origin water mass located in the upper mesopelagic layer (depth, 100–200 m). Below the halocline layer (depth, 300–3000 m), both the production and abundance decreased with depth, with log–log regression slopes of −1.33 and −0.77, respectively. Depth-integrated production and biomass in the meso- and bathy-pelagic layers was three- to five-fold lower than the corresponding values reported in the subpolar regions, whereas they were close to or lower than the corresponding values in oligotrophic subtropical regions. Prokaryote turnover times were estimated to be 1.1 and 6.1 years for meso- and bathy-pelagic layers, respectively, with the latter being among the longest turnover times reported for oceanic basins. We estimated prokaryote carbon demand in the water column (100–3000 m) to be on the order of 11 mg C m⁻² d⁻¹, which largely exceeds (by 38-fold) the sinking particulate organic carbon flux at depths of 120–200 m reported in the literature. This large carbon imbalance may be partly explained by organic carbon delivery by lateral intrusion of the Pacific-origin water mass into the upper mesopelagic layer.     

Organic biomarkers in the twilight zone—Time series and settling velocity sediment traps during MedFlux

The transfer of material through the twilight zone of the ocean is controlled by sinking particles that contain organic matter (OM) and mineral ballast. During the MedFlux field program in the northwestern Mediterranean Sea in 2003, sinking particulate matter was collected in time series (TS) and settling velocity (SV) traps and analyzed for amino acids, lipids, and pigments (along with ballast minerals) [Lee, C., Armstrong, R.A., Wakeham, S.G., Peterson, M.L., Miquel, J.C., Cochran, J.K., Fowler, S.W., Hirschberg, D., Beck, A. Xue, J., 2009b. Particulate matter fluxes in time series and settling velocity sediment traps in the northwestern Mediterranean Sea. Deep-Sea Research II, this volume [doi:10.1016/j.dsr2.2008.12.003]]. The goal was to identify how organic chemical compositions of sinking particles varied as a function of their in-situ settling velocity. The TS record was used to define the biogeochemical character and temporal pattern in flux during the period of SV trap deployment. Temporal variations in organic and mineral compositions are consistent with particle biogeochemistry being driven by the seasonal succession of phytoplankton. Spring diatom bloom conditions led to a high flux of rapidly sinking aggregates and zooplankton fecal matter; summer oligotrophy followed and was characterized by a higher proportion of slowly sinking phytoplankton cells. Bacterial degradation is particularly important during the low-flux summer period. Settling velocity traps show that a large proportion of particulate organic matter sinks at 200–500 m d⁻¹. Organic compositions of this fast-sinking material mirrors that of fecal pellets and aggregated material that sinks as the spring bloom terminates. More-slowly sinking OM bears a stronger signature of bacterial degradation than do the faster-sinking particles. The observation that compositions of SV-sorted fractions are different implies that the particle field is compositionally heterogeneous over a range of settling velocities. Thus physical and biological exchange between fast-sinking and slow-sinking particles as they pass down the water column must be incomplete.     

Relations between fecal pellet volume and body size for major zooplankters of the Inland Sea of Japan

Measurements of fecal pellet volume together with body length/body carbon weight were made for major zooplankters of the Inland Sea of Japan. The pellet volume was highly correlated with animal body size for copepods (10 species combined), a mysid (Neomysis japonica), a larvacean (Oikopleura dioica) and a pelagic shrimp (Acetes japonicus), and a specific equation was given for each group. A single equation could describe the composite relationship between pellet volume (PV, m³) and body carbon weight (C, g) for copepods andN. japonica: logPV=0.85logC+4.56. Balanid nauplii,O. dioica and a doliolidDolioletta gegenbauri produced pellets larger, butA. japonicus produced pellets smaller, than those by copepods andN. japonica of equivalent body carbon weight. In general, larger zooplankters produce larger fecal pellets. Hence, the size composition of the zooplankton community is an important parameter for the variation in the vertical flux of material via fecal pellets.