Diel variations of marine snow concentration in surface waters and implications for particle flux in the sea

Successive measurements of the size distribution and abundance of marine snow in the upper 100 m of the Santa Barbara Channel, California, with an in situ still camera system following 11 tagged water masses revealed a consistent pattern of nighttime decreases in the abundance of large particles. A net nocturnal reduction in particulate flux from the upper 100 m as calculated from camera profiles occurred in 75% of the day–night comparisons, and nighttime aggregate carbon losses resulted in a 38% average reduction in camera-derived aggregate flux. Intensive investigation of three stations for 24–48 h each indicated that nighttime decreases in aggregate concentrations and derived aggregate flux could be registered throughout the observed water column. Nocturnal decreases in marine snow concentration are unlikely to result from diel variations in the production of marine snow either as feeding webs of zooplankton or through variations in aggregation rates of smaller particles. Moreover, measured diel variations in the intensity of surface mixing and convective overturn during one of the 24 h deployments were not intense enough to produce aggregate fragmentation and reduced aggregate flux. Nighttime increases in large crustacean zooplankton (i.e., euphausiids and the large copepod Calanus pacificus) could explain some or all of the reduction in aggregate abundance at most stations. Fragmentation and consumption of marine snow by migrating macrozooplankton could produce our observed synchronous diel cycles in marine snow concentration. This is the first empirical evidence of a diel pattern in the concentration and calculated particulate flux of large sinking particles in near-surface waters. The data presented here are consistent with the only other existing diel study, which also reported decreases in marine snow abundance at night at 270 m depths in the oceanic north Atlantic. Diel variations in the sizes and concentrations of marine snow may impact water column processes dependent upon particle availability and size, such as grazing and remineralization, and may generate a diel cycle of food availability to the benthos.     

Suspended particle organic composition and cycling in surface and midwaters of the equatorial Pacific Ocean

In this study we relate spatial and temporal variation in the organic composition of suspended particles to current conceptual models of open-ocean particle cycling. Suspended particles in surface (0–200 m) and midwaters (200–1000 m) of the equatorial Pacific Ocean were collected during the 1992 US JGOFS Equatorial Pacific (EqPac) program. Samples collected during El Niño (Survey I) and normal conditions (Survey II) were analyzed for pigment, amino acid, fatty acid, and neutral lipid concentrations and compositions. Principal Components Analysis (PCA) and other statistical methods were used to assess changes in particulate organic composition between Surveys I and II, over 24° of latitude, from 15 to 850 m depth, and to compare our compositional data with previously published data from EqPac sinking particles. These analyses indicated that surface suspended particles (0–200 m) were similar in composition to surface ocean phytoplankton and were less degraded than particles sinking out of the euphotic zone (105 m). The organic composition of suspended particles in surface waters varied with latitudinal and El-Niño-induced changes in phytoplankton assemblages. Midwater suspended particles (200–1000 m) contained labile phytodetrital material derived from particles exiting the euphotic zone (105 m). However, labile organic constituents of midwater suspended particles were increasingly degraded by microbes or consumed by midwater metazoans with depth. The increase in degradation state observed for midwater suspended particles may also have been caused by dilution of deeper (450–850 m) suspended particle pools with more refractory material originating from fast-sinking particles, e.g., fecal pellets. However, the mechanism controlling midwater particle degradation state varied with flux regime; dilution of midwater suspended particles dominated only in the higher flux regime found at equatorial latitudes (5°N–5°S) during Survey II (normal conditions). In summary, it is apparent that organic matter alteration in midwaters, and not cycling within the euphotic zone, has the larger effect on organic composition of suspended particles in the deep equatorial Pacific Ocean.     

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.     

Fragmentation of marine snow by swimming macrozooplankton: A new process impacting carbon cycling in the sea

Comparisons of the abundances and size distributions of marine snow (aggregated particles >0.5 mm in diameter) in the upper 100 m of the water column at ten stations off Southern California in the late afternoon with those in the same parcel of water the following morning, after nocturnal vertical migration by zooplankton had occurred, revealed the existence of a previously undescribed process affecting marine particle dynamics. Aggregate abundances increased overnight and changes were positively and significantly correlated only with the abundance of the common euphausiid, Euphausia pacifica, and with no other biological or physical factor. Moreover, mean aggregate size decreased and aggregate size distributions shifted toward smaller size classes when euphausiids were abundant. The only conclusion consistent with these findings was that euphausiids were physically disaggregating marine snow into smaller, more numerous aggregates through shear stresses generated while swimming. Video-recording of both tethered and free-swimming E. pacifica in the laboratory dramatically confirmed that aggregates passing within 8–10 mm of the animal's abdomen were fragmented either by entrainment and direct impact with the beating pleopods or by eddies generated during swimming. At the abundances observed in this study, swimming E. pacifica would have sufficiently disturbed 3–33% of the water column each night to disrupt the aggregates contained therein. This is the first evidence for the fragmentation of large particles by the swimming activities of zooplankton and suggests that macrozooplankton and micronekton play a significant role in the particle dynamics of the water column regardless of whether they consume particles or not. Disaggregation of marine snow by swimming and migrating animals may alter the sizes of particles available to grazers and microbial colonizers and reduce the flux of particulate carbon by generating smaller particles, which potentially sink more slowly and reside longer in the water column. This newly discovered process reduces carbon flux while simultaneously conserving carbon and provides a previously unconsidered link between animal behavior and the biogeochemistry of the sea. It may help explain the exponential reduction in particle flux with depth observed in parts of the ocean and help balance oceanic carbon models.     

Seasonal and interannual variability in particle fluxes of carbon,nitrogen and silicon from time series of sediment traps at Ocean Station P, 1982–1993: relationship to changes in subarctic primary productivity

An extended time series of particle fluxes at 3800 m was recorded using automated sediment traps moored at Ocean Station Papa (OSP, 50°N, 145°W) in the northeast Pacific Ocean for more than a decade (1982–1993). Time-series observations at 200 and 1000 m, and short-term measurements using surface-tethered free-drifting sediment traps also were made intermittently. We present data for fluxes of total mass (dry weight), particulate organic carbon (POC), particulate organic nitrogen (PON), biogenic Si (BSi), and particulate inorganic carbon (PIC) in calcium carbonate. Mean monthly fluxes at 3800 m showed distinct seasonality with an annual minimum during winter months (December–March), and maximum during summer and fall (April–November). Fluxes of total mass, POC, PIC and BSi showed 4-, 10-, 7- and 5-fold increases between extreme months, respectively. Mean monthly fluxes of PIC often showed two plateaus, one in May–August dominated by <63 μm particles and one in October–November, which was mainly >63 μm particles. Dominant components of the mass flux throughout the year were CaCO₃ and opal in equal amounts. The mean annual fluxes at 3800 m were 32±9 g dry weight g m⁻² yr⁻¹, 1.1±0.5 g POC m⁻² yr⁻¹, 0.15±0.07 g PON m⁻² yr⁻¹, 5.9±2.0 g BSi m⁻² yr⁻¹ and 1.7±0.6 g PIC m⁻² yr⁻¹. These biogenic fluxes clearly decreased with depth, and increased during “warm” years (1983 and 1987) of the El Niño, Southern Oscillation cycle (ENSO). Enhancement of annual mass flux rates to 3800 m was 49% in 1983 and 36% in 1987 above the decadal average, and was especially rich in biogenic Si. Biological events allowed estimates of sinking rates of detritus that range from 175 to 300 m d⁻¹, and demonstrate that, during periods of high productivity, particles sink quickly to deep ocean with less loss of organic components. Average POC flux into the deep ocean approximated the “canonical” 1% of the surface primary production.     

Controls of 234Th removal from the oligotrophic ocean by polyuronic acids and modification by microbial activity

To gain new insights into the variability of particulate organic carbon (POC) fluxes and to better understand the factors controlling the POC/²³⁴Th ratios in suspended and sinking particulate matter, we investigated the relationships between POC/²³⁴Th ratios and biochemical composition (uronic acids, URA; total carbohydrates, TCHO; acid polysaccharides, APS; and POC) of suspended and sinking matter from the Gulf of Mexico in 2005 and 2006. Our data show that URA/POC in sediment traps (STs), APS/POC in the suspended particles, and turnover times of particulate ²³⁴Th in the water column and those of bacteria in STs inside eddies usually increased with depth, whereas particulate POC/²³⁴Th (10–50 μm) and the sediment-trap parameters (POC flux, POC/²³⁴Th ratio, bacterial biomass, and bacterial production) decreased with depth. However, this trend was not the case for most biological parameters (e.g., phytoplankton and bacterial biomass) or for the other parameters at the edges of eddies or at coastal-upwelling sites.In general, the following relationships were observed: 1) ²³⁴Th/POC ratios in STs were correlated with APS flux, and these ratios in the 10–50 μm suspended particles also correlated with URA/POC ratios; 2) neither URA fluxes nor URA/POC ratios were significantly related to bacterial biomass; 3) the sum of two uronic acids (G2, glucuronic, and galacturonic acid, which composed most of the URA pool) was positively related to bacterial biomass; and 4) the POC/²³⁴Th ratios in intermediate-sized particles (10–50 μm) were close to those in sinking particles but much lower than those in > 50 μm particles. The results indicate that acid polysaccharides, though a minor fraction (~ 1%) of the organic carbon, act more likely as proxy compound classes that might contain the more refractory ²³⁴Th-binding biopolymer, rather than acting as the original ²³⁴Th “scavenger” compound. Moreover, these acid polysaccharides, which might first be produced by phytoplankton and then modified by bacteria, also influence the on-and-off “piggy-back” processes of organic matter and ²³⁴Th, thus causing additional variability of the POC/²³⁴Th in particles of different sizes.     

Hydrolytic ectoenzyme activity associated with suspended and sinking organic particles within the anoxic Cariaco Basin

Ectohydrolase activities of suspended microbiota were compared to those associated with sinking particles (sed-POM) retrieved from sediment traps deployed in the permanently anoxic Cariaco Basin. In shore-based assays, activities of aminopeptidase, β-glucosidase, chitinase and alkaline phosphatase were measured in samples obtained from oxic and anoxic depths using MUF- and MCA-labeled fluorogenic substrate analogs. Hydrolysis potentials for these enzymes in the seston varied widely over the nine cruises sampled (8 Nov 1996–3 May 2000) and among depths (15–1265 m); from <10 to over 1600 nM d^?1 hydrolysate released, generally co-varying with one another and with suspended particulate organic carbon (POC) and particulate nitrogen (PN). Hydrolytic potentials, prokaryotic abundances and POC/PN concentrations in sinking debris were 400–1.3×10⁷ times higher than in comparable volumes of seawater. However when normalized to PN, hydrolytic potentials in sediment trap samples were not demonstrably higher than in Niskin bottle samples. We estimate that PN pools in sediment trap samples were turned over 2–1400 times (medians=7–26x) slower by hydrolysis than were suspended PN pools. Median prokaryotic growth rates (divisions d^?1) in sinking debris were also ～150 times slower than for bacterioplankton. Hydrolytic potentials in surface oxic waters were generally faster than in underlying anoxic waters on a volumetric basis (nM hydrolysate d^?1), but were not significantly (p>0.05) different when normalized to PN or prokaryote abundances. Alkaline phosphatase was consistently the most active ectohydrolase in both sample types, suggesting that Cariaco Basin assemblages were adapted to decomposing phosphate esters in organic polymers. However, phosphorus limitation was not evident from nutrient inventories in the water column. Results support the hypothesis that efficiencies of polymer hydrolysis in anoxic waters are not inherently lower than in oxic waters.     

Alkenones and particulate fluxes in sediment traps from the central equatorial Pacific

We investigated a year-long (September 1992 to August 1993) time series of total mass, calcium carbonate, organic carbon, opal, and alkenone fluxes in sinking particles collected with sediment traps moored at 1770 and 4220 m in the central equatorial Pacific. The total mass fluxes varied from 14.7 to 68.7 mg/m²/day at 1770 m, with greater fluxes in October–November and February–April, and from 14.6 to 50.4 mg/m²/day with peak fluxes during October–November at 4220 m. High flux in the spring season shown at 1770 m was not indicated at 4220 m; instead, a slight increase was shown during a broad period from March to June. The calcium carbonate fluxes varied from 10.8 to 49.1 mg/m²/day with higher fluxes in October–November and March–April at 1770 m, and from 8.9 to 37.0 mg/m²/day with a higher flux in October–November at 4220 m. The organic carbon fluxes varied from 0.36 to 5.91 mg/m²/day, with higher fluxes in October–November and March–April at 1770 m, and from 0.72 to 2.58 mg/m²/day at 4220 m. The annual mean organic carbon flux was 1.84 and 1.28 mg/m²/day at 1770 and at 4220 m, respectively. These values were less than half of those reported for the EqPac sediment trap experiment. The opal fluxes varied from 0.55 to 4.4 mg/m²/day at 1770 m and from 1.23 to 2.95 mg/m²/day at 4220 m. Alkenone fluxes varied significantly from 0.05 to 0.84 μg/m²/day, with high values in November, February–March, and June at 1770 m. For the 4220 m trap, these values ranged from 0.05 to 0.25 μg/m²/day, with slightly higher fluxes in April–May and June–July, which followed periods of high alkenone fluxes observed in February–April and June–July, respectively, at 1770 m depth. These values were remarkably low compared with those reported by the previous studies at other sites. U₃₇^K′ values were constantly high >0.95 throughout the collection period. However, relatively low U₃₇^K′ values (0.92 and 0.93) were occasionally observed during February to March. Estimated alkenone temperatures from those U₃₇^K′ values were about 27–29°C and consistent with the observed temperature of the upper layer at ca.100 m depth. The seasonal change of the U₃₇^K′ values could be affected by not only water temperature but also the relative amount of ‘warm’ and ‘cold’ types of alkenone producer in the central equatorial Pacific.     

Rapid lateral particle transport in the Argentine Basin: Molecular 14C and 230Thxs evidence

Recent studies have revealed that lateral transport and focusing of particles strongly influences the depositional patterns of organic matter in marine sediments. Transport can occur in the water column prior to initial deposition or following sediment re-suspension. In both cases, fine-grained particles and organic-rich aggregates are more susceptible to lateral transport than coarse-grained particles (e.g., foraminiferal tests) because of the slower sinking velocities of the former. This may lead to spatial and, in the case of redistribution of re-suspended sediments, temporal decoupling of organic matter from coarser sediment constituents. Prior studies from the Argentine Basin have yielded evidence that suspended particles are displaced significant distances (100–1000 km) northward and downslope by strong surface and/or bottom currents. These transport processes result in anomalously cold alkenone-derived sea-surface temperature (SST) estimates (up to 6 °C colder than measured SST) and in the presence of frustules of Antarctic diatom species in surface sediments from this area. Here we examine advective transport processes through combined measurements of compound-specific radiocarbon ages of marine phytoplankton-derived biomarkers (alkenones) from core tops and excess ²³⁰Th (²³⁰Th_xs)-derived focusing factors for late Holocene sediments from the Argentine Basin. On the continental slope, we observe ²³⁰Th_xs-based focusing factors of 1.4–3.2 at sites where alkenone-based SST estimates were 4–6 °C colder than measured values. In contrast, alkenone radiocarbon data suggest coeval deposition of marine biomarkers and planktic foraminifera, as alkenones in core tops were younger than, or similar in age to, foraminifera. We therefore infer that the transport processes leading to the lateral displacement of these sediment components are rapid, and hence probably occur in the upper water column (<1500 m).     

Episodic particulate fluxes at southern temperate mid-latitudes (42–45°S) in the Subtropical Front region,east of New Zealand

Sediment traps were deployed for almost 1 yr at two sites near 178°40′E in 1996–1997 on Chatham Rise (New Zealand). These sites were either side of the Subtropical Front (STF), which is a biologically productive zone, characterised by moderate atmospheric CO₂ uptake. At each site, PARFLUX sediment traps (Mk 7G–21) were deployed at 300 and 1000 m in 1500 m water depth. At 42°42′S, north of the STF, approximately 80% of the integrated total mass, POC and biogenic silica flux at 300 m occurred in a 7-day pulse in austral mid-spring (1064, 141 and 6 mg m⁻² d⁻¹, respectively, in early October). This pulse was recorded a week later in the 1000 m trap, indicating a particle sinking rate of 100 m d⁻¹. In contrast, at 44°37′S, south of the STF, the main flux of total mass and biogenic silica occurred 3 weeks later in late spring (289 and 3 mg m⁻² d⁻¹, respectively, in early November). Organic carbon, nitrogen and phosphorus fluxes were persistently high over spring at the southern site, although total POC flux integrated over 3 months was only 60 mg m⁻² d⁻¹. Thus, up to 2–3 times more material was exported north of the STF, compared with fluxes measured <200 km away to the south. As an integrated proportion of the annual total mass flux, however, more organic carbon was exported south of the STF (17% cf. 5–14%). Furthermore, organic material exported in spring from southern waters was labile and protein-rich (C : N — 8–16, C : P — 200–450, N : P — 13–36), compared to the more refractory, diatom-dominated material sinking out north of the STF in spring (C : N 9–22, C : P 50–230, N : P 5–19). These observations are consistent with anomalously high benthic biomass and diversity observed on south Chatham Rise. Resuspension and differential particle settling are probable causes for depth increases in particulate flux. Estimated particle source areas may be up to 120 km away due to high levels of mesoscale activity and mean flow in the STF region.     

Impact of lugworms (Arenicola marina) on mobilization and transport of fine particles and organic matter in marine sediments

We investigated the impact of sediment reworking fauna and hydrodynamics on mobilization and transport of organic matter and fine particles in marine sediments. Experiments were conducted in an annular flume using lugworms (Arenicola marina) as model organisms. The impact of lugworms on sediment characteristics and particle transport was followed through time in sediments experimentally enriched with fine particles (< 63 μm) and organic matter. Parallel experiments were run at low and high water current velocity (11 and 25 cm s^− 1) to evaluate the importance of sediment erosion at the sediment–water interface. There was no impact of fauna on sediment composition and particle transport at current velocity below the sediment erosion threshold. At current velocity above the erosion threshold, sediment reworking by lugworms resulted in dramatic particle transport (12 kg dry matter m^− 2) to an adjacent particle trap within 56 days. The transported matter was enriched 6–8 times in fine particles and organic matter when compared to the initial sediment. This study suggests that sediment reworking fauna is an important controlling factor for the particle composition of marine sediments. A. marina mediated sediment reworking greatly increases the sediment volume exposed to hydrodynamic forcing at the sediment–water interface, and through sediment resuspension control the content of fine particles and organic matter in the entire reworked sediment layer (> 20 cm depth).     

Spatial organization of particle size composition in an eddy–jet system off California

We describe here the results of an interdisciplinary study conducted off the coast of northern and central California during September 1993 in which we deployed an Optical Plankton Counter. This instrument counted and measured particles in the size range between 0.27 and 9.8 mm equivalent spheric diameter (ESD) occurring between the surface and 240 m depth. The survey region was characterized by the presence of the California Current jet and a cyclonic and an anticyclonic eddy. We analyzed the spatial (horizontal and vertical) distribution of planktonic particles and their relation to this hydrodynamic structure. We used specific analytical methods that take into account spatial constraints, i.e. autocorrelation analysis, constrained agglomerative clustering and contiguity constraints permutation analysis of variance. Horizontal spatial organization of particles was revealed at three different spatial scales (5, 18 and 100 km), while vertical patterns were described at a much smaller scale (20 m). We could detect some degree of similarity between particle size category spatial organization and hydrodynamic structure both by size category association independent of current movements and by comparison of dynamically differentiated areas. Five groups of similar size composition were detected that had some relation to the dynamic structure. Four sub-regions were determined a priori by their different hydrodynamic heights. We could describe a variability of particle abundance among these regions, both for total particles and for some size categories. Particles were more abundant inside the cyclonic eddy and less abundant inside the anticyclonic eddy. We also found deep concentration maxima inside the anticyclonic eddy and shallower concentration maxima inside the cyclonic eddy, with particles >2 mm ESD at deeper levels, for both daytime and nighttime sampling. No systematic difference was detected between daytime and nighttime samples in 0–240 m integrated total particle abundance. However, at night particles appeared to be concentrated into three depth strata (10–50, 70–90 and 90–230 m) of different size-abundance composition, while during the day particles were distributed into one shallow (10–50 m) and one deeper stratum (70–240 m). Smaller particles always occupied the most shallow depths     

Transformations of biogenic particulates from the pelagic to the deep ocean realm

This overview compares and contrasts trends in the magnitude of the downward Particulate Organic Carbon (POC) flux with observations on the vertical profiles of biogeochemical parameters in the NE subarctic Pacific. Samples were collected at Ocean Station Papa (OSP, 50°N, 145°W), between 18–22 May 1996, on pelagic stocks/rate processes, biogenic particle fluxes (drifting sediment traps, 100–1000 m), and vertical profiles of biogeochemical parameters from MULVFS (Multiple Unit Large Volume Filtration System) pumps (0–1000 m). Evidence from thorium disequilibria, along with observations on the relative partitioning of particles between the 1–53 μm and >53 μm classes in the 50 m mixed layer, indicate that there was little particle aggregation within the mixed layer, in contrast to the 50–100 m depth stratum where particle aggregation predominated. Vertical profiles of thorium/uranium also provided evidence of particle decomposition occuring at depths ca. 150 m; heterotrophic bacteria and mesozooplankton were likely responsible for most of this POC utilisation. A water column carbon balance indicated that the POC lost from sinking particles was the predominant source of carbon for bacteria, but was insufficient to meet their demands over the upper 1000 m. While, the vertical gradients of most parameters were greatest just below the mixed layer, there was evidence of sub-surface increases in microbial viability/growth rates at depths of 200–600 m. The C:N ratios of particles intercepted by free-drifting and deep-moored traps increased only slightly with depth, suggesting rapid sedimentation even though this region is dominated by small cells/grazers, and the upper water column is characterised by long particle residence times (>15 d), a fast turnover of POC (2 d) and a low but constant downward POC flux.     

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.     

A rapid birefringence method for measuring suspended CaCO3 concentrations in seawater

The extreme birefringence of calcium carbonate (CaCO₃) relative to other major components of marine particulate matter provides a basis for making optical in situ measurements of particulate inorganic carbon (PIC) in seawater. This concept was tested with a benchtop spectrophotometer equipped with a 1- and 10-cm path length sample cell and modified with linear polarizers to measure the birefringence of suspended particles. Sample suspensions containing 3–100% CaCO₃ (by weight) were prepared from calcareous marine sediment material and varying amounts of non-birefringent diatomaceous earth. The samples ranged in total suspended material from 0.003 to 249 mg l⁻¹ and PIC from 0.03 to 1820 μmol CaCO₃ l⁻¹. A positive relationship was observed between birefringence and PIC, with response falling off as the total particle concentration and the relative abundance of non-CaCO₃ particles in the sample increased. Sensitivity increased linearly with optical path length, and absolute detection limits of 0.2–0.4 and 0.04–0.08 μmol CaCO₃ l⁻¹, respectively, were determined for path lengths of 1- and 10-cm based on the intrinsic signal noise of the modified spectrophotometer. Conventional (i.e., non-polarized) transmittance measurements were used to correct the birefringence signal for the sensitivity loss due to interference from scattering and absorption. Without further modification, this spectrophotometer-based method can be used (with a 10-cm cell) to quantify PIC in most surface ocean waters—including those influenced by coccolithophore blooms. The spectrophotometer results define performance requirements and design parameters for an in situ instrument capable of operating over the oceanic range of PIC.     

Sea-ice-thickness variability in the Chukchi Sea,spring and summer 2002–2004

Measurements of sea-ice thickness were obtained from drill holes, an ice-based electromagnetic induction instrument (IEM), and a ship-borne electromagnetic induction instrument (SEM) during the early-melt season in the southern Chukchi Sea in 2002 and 2004, and in late summer 2003 at the time of minimum ice extent in the northern Chukchi Sea. An ice roughness criterion was applied to distinguish between level and rough or ridged ice. Ice-thickness modes in the probability density functions (PDFs) derived from drill-hole and IEM measurements agreed well, with modes at 1.5–1.6 and 1.8–1.9 m for all data from level ice. The PDFs derived from SEM measurements show that the primary modes are at 0.1 and 1.1 m in 2003 and 0.7 m in 2004. In 2002 and 2004, significant fractions (between one-third and one-half) of level ice were found to consist of rafted ice segments. Snow depth varied significantly between years, with 2004 data showing more than half the snow cover on level ice to be at or below 0.05 m depth in late spring. Ice growth simulations and examination of ice drift and deformation history indicate that impacts of atmospheric and oceanic warming on level-ice thickness in the region over the past few decades are masked to a large extent by variability in snow depth and the contribution of deformation processes. In comparison with submarine sonar ice-thickness data from previous decades, a reduction in ice thickness by about 0.5–1 m is in part explained by the replacement of multi-year with first-year ice over the Chukchi and Beaufort shelves.     

Vertical flux of particulate Al,Fe, Pb,and Ba from the upper ocean estimated from 234Th/238U disequilibria

The vertical sinking flux of particulate Al, Fe, Pb, and Ba from the upper 250 m of the Labrador Sea has been estimated from measurements of ²³⁴Th/²³⁸U disequilibrium and the respective metal/²³⁴Th ratios in >53 μm size particles. ²³⁴Th-derived particulate metal fluxes include in situ scavenged metals, labile lithogenic metals, and metals derived from external input (e.g., atmospheric supply). In contrast to the POC/²³⁴Th ratio, particle size-fractionated (0.4–10 μm, 10–53 μm, and >53 μm) Al/²³⁴Th, Fe/²³⁴Th and Pb/²³⁴Th, and Ba/²³⁴Th ratios generally increase with depth and exhibit no systematic change with particle diameter. Sinking fluxes of particulate Al (2.47–22.3 μmol m⁻² d⁻¹), Fe (2.69–16.3 μmol m⁻² d⁻¹), Pb (2.85–70 nmol m⁻² d⁻¹), and Ba (0.13–2.1 μmol m⁻² d⁻¹) at 50 m (base of the euphotic zone) and 100 m (base of the mixed layer) are largely within the range of previous sediment trap results from other ocean basins. Estimates of the upper ocean residence time of Al (0.07–0.28 yr) and Pb (0.8–2.9 yr) are short compared to previously reported values. The settling rate of >53 μm particles calculated from the ²³⁴Th data ranges from 14 to 38 m d⁻¹.     

Particulate organic carbon budget in the open Algero-Balearic Basin (Western Mediterranean): Assessment from a one-year sediment trap experiment

With the aim of improving the knowledge of the open ocean carbon cycle, we present a budget of particulate organic carbon (POC) fluxes carried out in the deep central part of the Algero-Balearic Basin (ABB) at 2850 m water depth based on a single mooring equipped with five automated sediment traps deployed from April 2001 to May 2002 at depths of 250, 845, 1440, 2145 and 2820 m. Suspended particulate matter (SPM) and superficial sediments were also used as indicators of hydrodynamics and carbon burial, respectively. The data reveal that the fraction of primary production buried in the sediment, which finally leads to the sequestration of carbon dioxide from the atmosphere, is 0.16%, lower than the values found in the nearby continental margin regions such as the Alboran Sea (0.48–0.89%) but of the same order as recorded at other Mediterranean sites at similar depths, such as the Ionian Sea (0.11%). As they sink through the water column, the particles exhibit decreases in flux that are similar to those observed elsewhere, but also show variations that appear to correlate with hydrological features of the water masses present in the basin, as revealed by SPM concentrations and compositions. The input of the tyrrhenian deep water (TDW) into the ABB at 800–1500 m of water depth exhibits low suspended POC concentrations and low sinking POC fluxes were also observed in this depth range. Gulf of Lions water mass formation appears to also contribute to elevated suspended POC concentrations and perhaps POC accumulation in the traps and sediments by spreading of dense cold water along the whole ABB that supplied POC at depths higher than 2000 m.     

Egg production of turbot,Scophthalmus maximus,in the Baltic Sea

In the brackish water Baltic Sea turbot spawn at ~ 6–9 psu along the coast and on offshore banks in ICES SD 24–29, with salinity influencing the reproductive success. The potential fecundity (the stock of vitellogenic oocytes in the pre-spawning ovary), egg size (diameter and dry weight of artificially fertilized 1-day-old eggs) and gonad dry weight were assessed for fish sampled in SD 25 and SD 28. Multiple regression analysis identified somatic weight, or total length in combination with Fulton's condition factor, as main predictors of fecundity and gonad dry weight with stage of maturity (oocyte packing density or leading cohort) as an additional predictor. For egg size, somatic weight was identified as main predictor while otolith weight (proxy for age) was an additional predictor. Univariate analysis using GLM revealed significantly higher fecundity and gonad dry weight for turbot from SD 28 (3378–3474 oocytes/g somatic weight) compared to those from SD 25 (2343 oocytes/g somatic weight), with no difference in egg size (1.05 ± 0.03 mm diameter and 46.8 ± 6.5 μg dry weight; mean ± sd). The difference in egg production matched egg survival probabilities in relation to salinity conditions suggesting selection for higher fecundity as a consequence of poorer reproductive success at lower salinities. This supports the hypothesis of higher size-specific fecundity towards the limit of the distribution of a species as an adaptation to harsher environmental conditions and lower offspring survival probabilities. Within SD 28 comparisons were made between two major fishing areas targeting spawning aggregations and a marine protected area without fishing. The outcome was inconclusive and is discussed with respect to potential fishery induced effects, effects of the salinity gradient, effects of specific year-classes, and effects of maturation status of sampled fish.     

Bacterivory in the northwestern Indian Ocean during the intermonsoon – northeast monsoon period

Bacterial grazing loss rates were studied by radioactive labeling of natural bacteria with L-(4,5-³H) leucine and from the rate of disappearance of bacterial cells in the northwestern Indian Ocean. Bacterivory was measured in a mixed sample that had been combined from various depths across the euphotic zone. Experiments were performed on 26 occasions at 19 stations in the Gulf of Oman and the Arabian Sea during the intermonsoon–northeast monsoon period (November–December 1994). Combined uptake of radiolabeled bacteria (ULB) in 1–8 and 8–100 μm size fractions was somewhat lower than loss of label (LBL) measured in the bacterial fraction (0.2–1.0 μm), suggesting loss of radioactivity from the grazers due to metabolism. The less sensitive rate of disappearance of bacterial cells (LBC) was on average 51% higher than LBL estimates. Results from ULB and LBL measurements revealed that bacterivory was higher in the Gulf of Oman (average loss rate 4.1% h^-1) than in the Arabian Sea where rates were slightly higher inshore (1.7% h^-1) than in the central gyre. Heterotrophic nanoflagellates in the 1–8 μm size fractions were identified as the primary bacterivores. Microzooplankton (8–100 μm) accounted for 33% of total bacterivory in the Gulf of Oman but only 16% in the central Arabian Sea. Time-course experiments conducted at two stations indicated that diel changes in bacterivory may be substantial in the northwestern Indian Ocean.