Investigating the effect of ballasting by CaCO3 in Emiliania huxleyi, II: Decomposition of particulate organic matter

The quantitative relationship between organic carbon and mineral contents of particles sinking below 1800 m in the ocean indicates that organisms with mineral shells such as coccolithophores are of special importance for transporting carbon into the deep sea. Several hypotheses about the mechanism behind this relationship between minerals and organic matter have been raised, such as mineral protection of organic matter or enhanced sinking rates through ballast addition. We examined organic matter decomposition of calcifying and non-calcifying Emiliania huxleyi cultures in an experiment that allowed aggregation and settling in rotating tanks. Biogenic components such as particulate carbon, particulate nitrogen, particulate volume, pigments, transparent exopolymer particles (TEP), and particulate amino acids in suspended particles and aggregates were followed over a period of 30 d. The overall pattern of decrease in organic matter, the amount of recalcitrant organic matter left after 30 d, and the compositional changes within particulate organic matter indicated that cells without a shell are more subject to loss than calcified cells. It is suggested that biogenic calcite helps in the preservation of particulate organic matter (POM) by offering structural support for organic molecules. Over the course of the experiment, half the particulate organic carbon in both calcifying and non-calcifying cultures was partitioned into aggregates and remained so until the end of the experiment. The partial protection of particulate organic matter from solubilization by biominerals and by aggregation that was observed in our experiment may help explain the robustness of the relationship between organic and mineral matter fluxes in the deep ocean.     

The distribution and flux of particulate matter in the Bideford River Estuary,Prince Edward Island,Canada

The temporal and spatial distribution of total and organic particulate matter is investigated in the Bideford River estuary. Particulate matter is homogenously distributed in both the water column and the surface sediment, due to high rates of resuspension and lateral transport. The measured mean sedimentation rate for the estuary is 183·5 g of particulate matter m^?2 day^?1, of which more than half is due to resuspension.The surface sediment of the estuary is quantitatively the dominant reservoir of organic matter, with an average of 902·5 g of particulate organic carbon (POC) m^?2 and 119·5 g of particulate organic nitrogen (PON) m^?2. Per unit surface area, the sediment contains 450 times more POC and 400 times more PON than the water column. Terrestrial erosion contributes high levels of particulate matter, both organic and inorganic, to the estuary from the surrounding watershed. Low rates of sediment export from the estuary result in the accumulation of the terrigenous material. The allochthonous input of terrigenous organic matter masks any relationship between the indigenous plant biomass and the organic matter.In the water column, a direct correlation exists between the organic matter, i.e. POC and PON, concentration and the phytoplankton biomass as measured by the plant pigments. Resuspension is responsible for the residual organic matter in the water column unaccounted for by the phytoplankton biomass.The particulate content of the water column and the surface sediment of the estuary is compared to that of the adjacent bay. Water-borne particulate matter is exported from the estuary to the bay, so that no significant differences in concentration are noted. The estuarine sediment, however, is five to six times richer in organic and silt-clay content than the bay sediment. Since sediment flux out of the estuary is restricted, the allochthonous contribution of terrigenous particulate matter to the bay sediment is minor, and the organic content of the bay sediment is directly correlated to the autochthonous plant biomass.     

Relationship between the suspended particulate matter and microorganisms in the White Sea waters

During the summer periods of 2003–2005, the spatial and vertical distributions of the suspended particulate matter and bacteria in stratified and homogeneous waters of the White Sea was studied. The results of the study of various quantitative characteristics of the suspended particulate matter (first of all, the area of the surface and the volumetric and mass concentrations) and the abundance of microorganisms in the water are discussed. A direct correlation between the value of the surface area of the suspended particulate matter and the total number of bacteria in the water is revealed. However, it was manifested only during the early summer period of the observations and was not expressed at the end of the summer. The enhanced surface area of the suspended particulate matter can indicate the higher biochemical activity of its particles. The influence of the pelitic fraction on the bacteria abundance in different parts of the sea during the summer is estimated.     

Properties of coarse particles in suspended particulate matter of the North Yellow Sea during summer

Fine particles in seawater commonly form large porous aggregates. Aggregate density and settling velocity determine the behavior of this suspended particulate matter(SPM) within the water column.However, few studies of aggregate particles over a continental shelf have been undertaken. In our case study, properties of aggregate particles, including size and composition, over the continental shelf of the North Yellow Sea were investigated. During a scienti?c cruise in July 2016, in situ ef fective particle size distributions of SPM at 10 stations were measured, while temperature and turbidity measurements and samples of water were obtained from surface, middle, and bottom layers. Dispersed and inorganic particle size distributions were determined in the laboratory. The in situ SPM was divided into(1) small particles(<32 μm),(2) medium particles(32–256 μm) and(3) large particles(>256 μm). Large particles and medium particles dominated the total volume concentrations(VCs) of in situ SPM. After dispersion, the VCs of medium particles decreased to low values(<0.1 μL/L). The VCs of large particles in the surface and middle layers also decreased markedly, although they had higher peak values(0.1–1 μL/L). This suggests that almost all in situ medium particles and some large particles were aggregated, while other large particles were single particles. Correlation analysis showed that primary particles <32 μm in?uenced the formation of these aggregates. Microscopic examination revealed that these aggregates consisted of both organic and inorganic ?ne particles, while large particles were mucus-bound organic aggregates or individual plankton.The vertical distribution of coarser particles was clearly related to water strati?cation. Generally, medium aggregate particles were dominant in SPM of the bottom layer. A thermocline blocked resuspension of?ne material into upper layers, yielding low VCs of medium-sized aggregate particles in the surface layer.Abundant large biogenic particles were present in both surface and middle layers.     

Vertical Patterns of Suspended Matter Characteristics Along a Coastal-basin Transect in the Western Baltic Sea

Suspended particulate matter samples were collected from the water column, the bottom nepheloid layer and the ‘ fluffy layer ’ from four stations along a coastal-basin transect in the Pomeranian Bight, western Baltic Sea. Sampling was performed nine times between October 1996 and December 1998 for various analyses, including electron probe x-ray micro analysis for detailed mineralogical investigations.Specific vertical patterns of clay mineral distributions were found. Suspended particulate matter (SPM) in the bottom nepheloid layer and the ‘ fluffy layer ’ overlying sediments was enriched in organic carbon and hydrated three layer clay minerals, whereas the non-aggregated SPM was dominated by quartz and biogenic opal. It appears that separation effects operate during aggregation of mineral particles and organic matter in repeated cycles of resuspension and settling. No clear seasonal variations in the composition of the SPM were found, in spite of high spatial and temporal variability of biological and physical variables. The results suggest that preferential incorporation, possibly aided by microbiological colonization, of hydrated three layer silicates into the organic flocs is a process that occurs under a wide range of conditions. Because aggregates sink faster than individual particles, aggregate formation led to a relative enrichment of illite and smectite in the near-bottom layers. Considering the affinity of organic contaminants and heavy metals to organic matter, the selective removal of aggregated organic matter and hydrated three-layer clay minerals from the water column and enhanced transport in the near-bottom fluffy layer may be a natural cleansing mechanism operating in the shallow waters of the bight.     

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.     

A mesocosm experiment coupled with optical measurements to assess the fate and sinking of atmospheric particles in clear oligotrophic waters

It has recently been postulated that lithogenic particles such as Saharan dust strongly influence particulate organic carbon export to the deep ocean by acting as mineral ballast. However, our understanding of the processes involved remains scant. In the present study, optical measurements were performed to monitor variations in the concentration, composition and size distribution of particles in suspension within the water column after simulating a Saharan dust event in very clear Mediterranean waters off Corsica in June 2010. A new methodology set up in large mesocosms proved very successful in this regard. Values obtained simultaneously from three instruments (WetLabs ECO-BB3, WetLabs ac-9, Sequoia Scientific LISST-100) provided evidence that (1) part of the Saharan dust pool has a rapid settling velocity (∼24–86 m day⁻¹), (2) particulate export following a dust event is a nonlinear multi-step process and (3) export is controlled in part by the formation of organic-mineral aggregates. This experimental study provides the first insight of the complex export processes occurring after a dust event involving both physical and biogeochemical forcings in clear oligotrophic waters.     

Manganese recycling in coastal waters

Both water and suspended particulate matter in the deep water of the Gulf of St. Lawrence are greatly enriched in manganese. Maximum dissolved manganese concentrations are encountered close to the sediment-water interface whereas the particulate matter with highest manganese content occurs 30–100 m above the bottom. The elevated concentrations are attributed to the diagenetic release of dissolved manganese from the underlying fine-grained sediments and its subsequent precipitation in the water column. The rate of manganese precipitation is rapid compared to the rates of diffusion and mixing in the bottom water. Part of the manganese-enriched particulate matter becomes mixed throughout the water column by advection and diffusion. Thus, particles enriched in manganese can ultimately be carried into the open ocean by prevailing currents. This process, which appears to be widespread in eastern Canadian coastal waters, enables manganese originally associated with rapidly settling terrigenous particles to be transferred to slowly settling fine-grained suspended particles entering the ocean from coastal environments. In this way, riverborne manganese of terrigenous origin may well account for a major proportion of the excess manganese in pelagic sediments.     

Particulate barium fluxes on the continental margin: a study from the Alboran Sea (Western Mediterranean)

Particulate biogenic barium (bio-Ba) fluxes obtained from three instrumented arrays moored in the Alboran Sea, the westernmost basin in the Mediterranean Sea, are presented in this study. The mooring lines were deployed over almost 1 year, from July 1997 to May 1998, and were equipped with sediment traps at 500–700 m depth, 1000–1200 m depth and 30 m above the seafloor (1000–2200 m). The results obtained support the growing body of evidence that the relationship between particulate bio-Ba and Corg throughout the water column in margin systems is clearly different from this relation in the open ocean. In the Alboran Sea, the annual averaged bio-Ba fluxes range from 0.39 to 1.07 μmol m⁻² day⁻¹, with mean concentrations of 1.31–1.69 μmol g⁻¹ and bio-Ba/Corg ratios lower than in the open ocean. The low bio-Ba values obtained also indicate that calculating bio-Ba is extremely sensitive to the detrital Ba/Al ratio of each sample. The lithogenic Ba fraction in the Alboran Sea continental margin area contributes between 24% and 85% of the total Ba. Increased bio-Ba export efficiency was observed after periods of high primary productivity and suggests that the processes limiting the bio-Ba formation in the study area relate to settling dynamics of organic matter aggregates. Furthermore, the ballasting effect of the abundant lithogenic and carbonate particles may limit decomposition of organic matter aggregates and enhance the transfer of particles rich in Corg and relatively poor in bio-Ba to the deep seafloor. Lateral input of freshly sedimented biogenic material, including particulate bio-Ba, has been observed on the lower continental slope in the western Alboran Sea. These observations emphasize that the use of the bio-Ba as a proxy of export productivity from the surface ocean must be used cautiously in highly dynamic environments such as those in the Alboran Sea.     

Factors controlling aggregate formation in the benthic boundary layer of the Mecklenburg Bight (western Baltic Sea)

Studies on aggregate formation and size distribution in relation to bottom water composition and flow regime were carried out in November 1994 at two transects in the inner and outer Mecklenburg Bight (Baltic Sea). The bottom water sampler ‘BIOPROBE' (BWS) was used to collect 10-dm³ water samples at 5, 10, 20 and 40 cm above the seabed. The outer transect samples tended to be more influenced by the open western Baltic Sea, whereas the inner transect samples were more affected by the coastal hydrography. Aggregate size distribution was investigated using a newly developed particle camera allowing identification of particles down to 150 μm size. Increasing concentrations of total particulate matter (TPM), particulate organic carbon (POC) and chlorophyll pigment equivalents (CPE) towards the seafloor together with a low proportion of POC/TPM (<5%) implied that the material was of resuspended origin. Aggregate size in both transects was positively correlated with TPM, transparent exopolymer particles (TEP) and bacterial cell abundance. Higher particle concentrations and aggregate numbers in the outer transect indicated a higher resuspension frequency, or lateral advection processes. The higher concentration of aggregates at the outer transect may reflect the larger amount of near-bottom transported material.     

Significance of the Virus-Rich 2–200 nm Size Fraction of Seawater for Heterotrophic Flagellates: I. Impact on Growth

Abstract. The effect of artificially changing the concentration of naturally occurring free virus-like particles (VLPs) on growth of marine heterotrophic flagellates was tested in batch cultures using ultrafiltration technology. This manipulation influenced the course of flagellate growth markedly. During 1 week of incubation the growth of 4 tested flagellate strains ( Oxyrrhis marina, Paraphysomonas imperforata, Petalomonas cantuscygni, Pteridomonas danica ) was strongly favoured after increasing the VLP concentration. Bacteria, however, were repressed in these treatments beyond 50 h. In cultures with natural seawater microbiota, we both reduced and enriched the concentration of VLPs. In these cultures, reduction of material in the virus-rich 2–200 nm size fraction led to a strong positive growth response of heterotrophic flagellates. In VLP-enriched treatments of natural seawater microbiota a tendency toward growth stimulation was also found, although this was not significant. Enrichment with VLPs caused no recognizable mortality either in cultured flagellate strains or in naturally occurring flagellate communities. However, it is suggested that a highly bioactive component must be present in the virus-rich 2–200 nm size fraction. Virus-like particles are discussed as possible candidates influencing heterotrophic flagellate community successions.     

Temporal variations of dissolved and particulate Th at a coastal station of the northern Adriatic Sea

Water–particle interactions, particle behaviour and short-time scale variability were assessed at a coastal station adjacent to the Emilia Romagna Region (Adriatic Sea) using dissolved and particulate ²³⁴Th analyses. The water column was sampled six times between March and September 1997. Measurements showed that ²³⁴Th is actively scavenged by particles but the dissolved fraction is always prevalent. Changes in hydrological conditions affect to some degree thorium activities and residence times. Dissolved thorium inventories slowly increased from May to July, then decreased in August, and increased again in September. In July, the formation of a sharp pycnocline associated with low productivity led to high dissolved and very low particulate ²³⁴Th activities due to inefficient scavenging. The presence of mucilaginous aggregates, observed in both August and September, may have played a role in scavenging of thorium. In September at 16 m depth, the highest ²³⁴Th particulate activity of the study period was measured, probably due to the presence of mucilage. However, the thorium deficit was scarce, due to the small sinking velocity of these aggregates. Both steady-state and non-steady state models were used to calculate residence times for the whole water column and its topmost part (10 m) obtaining strictly comparable results. Residence times in the whole water column are small, ranging from 15 to 45 and from 0.5 to 24 d for dissolved and particulate thorium, respectively.     

溶解有机碳在混合水中的行为研究

研究溶解有机碳(DOC)在海水和黄河水所组成的混合水中的迁移转化行为,结果表明,海水和已过滤黄河水混合时,DOC呈近似的保守性;海水与未过滤的黄河水混合时,DOC呈明显的非保守性。通过研究沉积物对DOC的解吸过程,发现沉积物在盐水中能解吸出DOC,随着盐度的增大,其解吸量也迅速增大。因此可以认为,DOC在河口区的非保守行为是DOC的絮凝和颗粒物的解吸共同作用的结果。     

High molecular weight organic matter in seawater

The presence and nature of high molecular weight organic matter in seawater was critically reviewed and its biogeochemical cycle was discussed.Organic matter that passes through a filter of 0.5–1 μm pore size is called dissolved and that which does not pass through such a filter is defined as particulate. The size of colloidal particles ranges from 0.001 to 1 μm, and therefore, they are included in the dissolved fraction having high molecular weight.High molecular weight organic matter greater than 100,000 molecular weight was found in the seawater of Tokyo Bay. The values ranged from 0.1 to 1.5 mgC/l, and accounted for 8–45% of the total dissolved organic matter.Decomposition experiments on dissolved organic matter showed that macromolecular organic matter is refractory to bacterial attack. However, macromolecular organic materials tend to aggregate or adsorb on small particles to a sufficient size for precipitation. Organic aggregates thus formed sink to the bottom of the sea and bioelements included in them are removed from water column. High molecular weight organic materials are, therefore, considered to play an important role in transportation and distribution of matter in seawater.In order to elucidate the chemical and biological properties of macromolecular organic matter, concentration and isolation of this material are important, using methods such as adsorption on organic adsorbents or ultrafiltration.     

Sedimentary matter fluxes in the contour current sedimentation system over the continental slope of the Norwegian Sea

The sedimentation system of the bottom contour current over the continental slope of Bear Island in the Norwegian Sea is considered. The nepheloid layer that provides the high horizontal flux of sedimentary material represents the main source of matter for the bottom sediments. The vertical particulate matter flux is largely formed in the nepheloid layer; the flux from higher layers of the water column is insignificant. Horizontal and vertical fluxes of sedimentary matter show a positive correlation. The flux of the matter from the bottom sediment into the nepheloid matter and the residence time of particles in the latter are estimated.     

A review of the effects of particle types on oil-suspended particulate matter aggregate formation

Oil-suspended particulate matter aggregate (OSA) can form naturally when oil and particles interact. The interaction between oil and suspended particulate matter makes oil less sticky, and facilitates its dispersion in the water column. The high oil-water surface contact enhances the biodegradation of oil and thus increases the efficiency of remediation processes. There are many factors that affect OSA formation, but, particle type is one of the most important. Because different particle types have different physical, chemical, and biological properties, their interactions with oil differ greatly. Particle properties such as interlayer spaces, hydrophobicity, surface charges, polarity, organic content, and size affect the interactions between materials and oil. These different interactions determine the type, buoyancy, size, and stability of OSA that forms, thus determining its fate in the environment. This review provides a current understanding of (1) OSA formation mechanisms, (2) sources and classes of marine materials, (3) oil-particle interactions, (4) material properties and their effects on oil interaction, and (5) future research needs.     

Rare earth elements in seawater during an iron-induced phytoplankton bloom of the western subarctic Pacific (SEEDS-II)

During an iron-enrichment experiment in the western subarctic Pacific (SEEDS-II), concentrations of dissolved and acid-soluble rare earth elements (REEs) were determined in shallow waters. Detailed vertical profiles of dissolved REEs were compared with those of nutrients in a preliminary survey. The results showed good correlation, except for Ce and Y. Along with phytoplankton growth within the iron-enriched patch, dissolved REEs were decreased in surface waters. The changes in REE concentrations were small but systematic: light REEs were adsorbed preferentially over heavy REEs. This systematic trend is consistent with results of earlier studies of REE adsorption onto biogenic particle surfaces. However, we were unable to detect a decrease of acid-soluble REEs in surface waters. During the phytoplankton bloom decline phase, dissolved REEs were elevated to concentration levels at the preliminary survey, although nutrients were decreased. The liberation of REEs from the suspended particles might be related to disaggregation from large particles to colloid particles, desorption from the particulate phase, and remineralization from organic matter. Outside the Fe-patch, time variations of REEs were also observed during SEEDS-II, indicating that both artificially induced phytoplankton blooms and natural variation of biogenic particles can affect the biogeochemical cycles of REEs in shallow waters within a short time.     

The biogeochemical fate and toxicity of mercury in Controlled Experimental Ecosystems

The affinity of mercury for organic matter was the most important parameter governing its chemical speciation, transport and toxicity in Controlled Experimental Ecosystems (CEEs) of the Controlled Ecosystem Pollution Experiment (CEPEX) deployed in Saanich Inlet, B.C., Canada, Particulate, colloidal and high molecular weight dissolved forms of mercury accounted for about 90% of the total mercury present in the water column. Bioassays indicated that organic matter in the above size classes had an important influence on the toxicity of mercury to phytoplankton.Sorption of mercury to particulate organic matter in the CEEs could be described by either Freundlich or linear adsorption isotherms. An observed depth-dependent increase in mercury content of particulate matter was attributed to selective biological degradation of non-mercury-binding organic components of the particles during settling.The high affinity of mercury for settling particulate organic matter led to rapid removal of mercury from CEEs. Removal rates could be described as first order and were a function of primary production rates. Deviations of measured half-removal times from those predicted by an empirically formulated relationship between primary production and mercury half-removal times could be related successfully to the influence of other occasionally significant biological parameters such as zooplankton grazing activity.     

Distribution of 210Po and export of organic carbon from the euphotic zone in the southwestern East Sea (Sea of Japan)

The distribution of the natural radionuclide ²¹⁰Po in the water column along a horizontal transect of the continental shelf, slope and deep basin regions of the East Sea (Sea of Japan), a marginal sea of the Northwest Pacific Ocean, was investigated, and its behavior is described here. The settling fluxes of particulate ²¹⁰Po in the deep basin along with ²¹⁰Pb, ²³⁴Th and biogenic matter were also determined. ²¹⁰Po inventories in the water column were observed to decrease from winter to summer in all stations, probably due to increased influx of ²¹⁰Po-poor Kuroshio Water of the Northwest Pacific Ocean during summer. Vertical profiles of dissolved and particulate ²¹⁰Po along with the settling fluxes of particulate ²¹⁰Po in the deep basin station have enabled us to evaluate temporal variations and residence times of ²¹⁰Po. In the slope and basin, activities of dissolved ²¹⁰Po generally decreased from the surface to the bottom water, with maximum activity just below the subsurface chlorophyll a maximum at 50–75 m depth in spring and summer. These subsurface peaks of dissolved ²¹⁰Po activity were attributed to the release of ²¹⁰Po from the decomposition of ²¹⁰Po-laden biogenic particulate organic matter. In the deep basin, despite the decrease in total mass flux, the sinking flux of particulate ²¹⁰Po was higher in the deeper trap (2000 m) than in the shallower one (1000 m), probably due to scavenging of dissolved ²¹⁰Po from the water column during particle descent and/or break-down of ²¹⁰Po-depleted particulate matter between 1,000 m and 2,000 m depths. In general, the ratios of the particulate phase to the dissolved phase of ²¹⁰Po (K_d) increased with depth in the slope and basin stations. ²¹⁰Po removal from the water column appears to depend on the primary productivity in the upper waters. There is an inverse relationship between K_d and suspended particulate matter (SPM) concentration in the water column. From the ²¹⁰Po activity/chlorophyll a concentration ratios, it appears that sinking particles arriving at 1000 m depth were similar to those in the surface waters.     

Particle flux across the mid-European continental margin

Results are presented from particle flux studies using sediment trap and current meter moorings along a transect at the European continental margin at 49°N within the EU-funded Ocean Margin Exchange (OMEX) project. Two moorings were placed, at the mid- and outer slope in water depths of 1500 and 3660 m, with traps at 600 and 1050 m and at 580, 1440 and 3220 m, respectively. Residual currents at the mid-slope follow the slope contour, whereas seasonal off-slope flow was registered at the outer slope. At 600 m on the slope fluxes are similar to those in the abyssal North Atlantic. The flux of all components (bulk dry weight, particulate organic and inorganic carbon, lithogenic matter and opal) increased with water depth. Highest fluxes were recorded at 1440 m at the outer slope, where off-slope residual currents mediate particle export. The injection of biogenic and lithogenic particles below the depth of winter mixing results in the export of particles from shallower waters. Calculated lateral fluxes of particulate organic carbon exceed the primary flux by over a factor of 2 at 1440 m on the outer slope. Estimated lateral fluxes of suspended particulate matter in the water column and intermediate nepheloid layers at the outer slope are potentially large compared to sinking fluxes measured by sediment traps. A comparison is made of particle flux at three continental margin sites and two sites in the adjacent open North Atlantic, from which it is seen that bulk and organic matter flux increases exponentially with proximity to the shelf break. The percentage contribution of particulate organic carbon to biogenic fluxes increases from a mean of 5.7% in the abyssal N. Atlantic to 13.9% at the continental margins.