Food web flows through a sub-arctic deep-sea benthic community

The benthic food web of the deep Faroe–Shetland Channel (FSC) was modelled by using the linear inverse modelling methodology. The reconstruction of carbon pathways by inverse analysis was based on benthic oxygen uptake rates, biomass data and transfer of labile carbon through the food web as revealed by a pulse-chase experiment. Carbon deposition was estimated at 2.2 mmol C m⁻² d⁻¹. Approximately 69% of the deposited carbon was respired by the benthic community with bacteria being responsible for 70% of the total respiration. The major fraction of the labile detritus flux was recycled within the microbial loop leaving merely 2% of the deposited labile phytodetritus available for metazoan consumption. Bacteria assimilated carbon at high efficiency (0.55) but only 24% of bacterial production was grazed by metazoans; the remaining returned to the dissolved organic matter pool due to viral lysis. Refractory detritus was the basal food resource for nematodes covering ∼99% of their carbon requirements. On the contrary, macrofauna seemed to obtain the major part of their metabolic needs from bacteria (49% of macrofaunal consumption). Labile detritus transfer was well-constrained, based on the data from the pulse-chase experiment, but appeared to be of limited importance to the diet of the examined benthic organisms (<1% and 5% of carbon requirements of nematodes and macrofauna respectively). Predation on nematodes was generally low with the exception of sub-surface deposit-feeding polychaetes that obtained 35% of their energy requirements from nematode ingestion. Carnivorous polychaetes also covered 35% of their carbon demand through predation although the preferred prey, in this case, was other macrofaunal animals rather than nematodes. Bacteria and detritus contributed 53% and 12% to the total carbon ingestion of carnivorous polychaetes suggesting a high degree of omnivory among higher consumers in the FSC benthic food web. Overall, this study provided a unique insight into the functioning of a deep-sea benthic community and demonstrated how conventional data can be exploited further when combined with state-of-the-art modelling approaches.     

Comparative biomass structure and estimated carbon flow in food webs in the deep Gulf of Mexico

A budget of the standing stocks and cycling of organic carbon associated with the sea floor has been generated for seven sites across a 3-km depth gradient in the NE Gulf of Mexico, based on a series of reports by co-authors on specific biotic groups or processes. The standing stocks measured at each site were bacteria, Foraminifera, metazoan meiofauna, macrofauna, invertebrate megafauna, and demersal fishes. Sediment community oxygen consumption (SCOC) by the sediment-dwelling organisms was measured at each site using a remotely deployed benthic lander, profiles of oxygen concentration in the sediment pore water of recovered cores and ship-board core incubations. The long-term incorporation and burial of organic carbon into the sediments has been estimated using profiles of a combination of stable and radiocarbon isotopes. The total stock estimates, carbon burial, and the SCOC allowed estimates of living and detrital carbon residence time within the sediments, illustrating that the total biota turns over on time scales of months on the upper continental slope but this is extended to years on the abyssal plain at 3.6 km depth. The detrital carbon turnover is many times longer, however, over the same depths. A composite carbon budget illustrates that total carbon biomass and associated fluxes declined precipitously with increasing depth. Imbalances in the carbon budgets suggest that organic detritus is exported from the upper continental slope to greater depths offshore.The respiration of each individual “size” or functional group within the community has been estimated from allometric models, supplemented by direct measurements in the laboratory. The respiration and standing stocks were incorporated into budgets of carbon flow through and between the different size groups in hypothetical food webs. The decline in stocks and respiration with depth were more abrupt in the larger forms (fishes and megafauna), resulting in an increase in the relative predominance of smaller sizes (bacteria and meiofauna) at depth. Rates and stocks in the deep northern GoM appeared to be comparable to other continental margins where similar comparisons have been made.     

Carbon flows in the benthic food web at the deep-sea observatory HAUSGARTEN (Fram Strait)

The HAUSGARTEN observatory is located in the eastern Fram Strait (Arctic Ocean) and used as long-term monitoring site to follow changes in the Arctic benthic ecosystem. Linear inverse modelling was applied to decipher carbon flows among the compartments of the benthic food web at the central HAUSGARTEN station (2500 m) based on an empirical data set consisting of data on biomass, prokaryote production, total carbon deposition and community respiration. The model resolved 99 carbon flows among 4 abiotic and 10 biotic compartments, ranging from prokaryotes up to megafauna. Total carbon input was 3.78±0.31 mmol C m⁻² d⁻¹, which is a comparatively small fraction of total primary production in the area. The community respiration of 3.26±0.20 mmol C m⁻² d⁻¹ is dominated by prokaryotes (93%) and has lower contributions from surface-deposit feeding macro- (1.7%) and suspension feeding megafauna (1.9%), whereas contributions from nematode and other macro- and megabenthic compartments were limited to <1%. The high prokaryotic contribution to carbon processing suggests that functioning of the benthic food web at the central HAUSGARTEN station is comparable to abyssal plain sediments that are characterised by strong energy limitation. Faunal diet compositions suggest that labile detritus is important for deposit-feeding nematodes (24% of their diet) and surface-deposit feeding macrofauna (∼44%), but that semi-labile detritus is more important in the diets of deposit-feeding macro- and megafauna. Dependency indices on these food sources were also calculated as these integrate direct (i.e. direct grazing and predator–prey interactions) and indirect (i.e. longer loops in the food web) pathways in the food web. Projected sea-ice retreats for the Arctic Ocean typically anticipate a decrease in the labile detritus flux to the already food-limited benthic food web. The dependency indices indicate that faunal compartments depend similarly on labile and semi-labile detritus, which suggests that the benthic biota may be more sensitive to changes in labile detritus inputs than when assessed from diet composition alone. Species-specific responses to different types of labile detritus inputs, e.g. pelagic algae versus sympagic algae, however, are presently unknown and are needed to assess the vulnerability of individual components of the benthic food web.     

Mesopelagic zone ecology and biogeochemistry – a synthesis

The mesopelagic zone is the oceanic region through which carbon and other elements must pass in order to reach deeper waters or the sea floor. However, the food web interactions that occur in the mesopelagic zone are difficult to measure and so, despite their crucial importance to global elemental cycles, are not very well known. Recent developments in technology and new approaches have advanced the study of the variability in and controls upon the distribution and diversity of organisms in the mesopelagic zone, including the roles of respiration, recycling, and repackaging of particulate and dissolved organic material. However, there are remarkably few syntheses of the ecology and biogeochemistry of the microbes and metazoa that permanently reside or habitually visit this ‘twilight zone’. Without this synthesis, it is difficult to assess the impact of ongoing changes in ocean hydrography and chemistry, due to increasing atmospheric carbon dioxide levels, on the biological carbon pump. This paper reviews what is known about the distribution of microbes and metazoa in the mesopelagic zone in relation to their activity and impact on global biogeochemical cycles. Thus, gaps in our knowledge are identified and suggestions made for priority research programmes that will improve our ability to predict the effects of climate change on carbon sequestration.     

Feeding ecology of mesopelagic zooplankton of the subtropical and subarctic North Pacific Ocean determined with fatty acid biomarkers

Mesopelagic zooplankton may meet their nutritional and metabolic requirements in a number of ways including consumption of sinking particles, carnivory, and vertical migration. How these feeding modes change with depth or location, however, is poorly known. We analyzed fatty acid (FA) profiles to characterize zooplankton diet and large particle (>51 μm) composition in the mesopelagic zone (base of euphotic zone ?1000 m) at two contrasting time-series sites in the subarctic (station K2) and subtropical (station ALOHA) Pacific Ocean. Total FA concentration was 15.5 times higher in zooplankton tissue at K2, largely due to FA storage by seasonal vertical migrators such as Neocalanus and Eucalanus. FA biomarkers specific to herbivory implied a higher plant-derived food source at mesotrophic K2 than at oligotrophic ALOHA. Zooplankton FA biomarkers specific to dinoflagellates and diatoms indicated that diatoms, and to a lesser extent, dinoflagellates were important food sources at K2. At ALOHA, dinoflagellate FAs were more prominent. Bacteria-specific FA biomarkers in zooplankton tissue were used as an indicator of particle feeding, and peaks were recorded at depths where known particle feeders were present at ALOHA (e.g., ostracods at 100–300 m). In contrast, depth profiles of bacterial FA were relatively constant with depth at K2. Diatom, dinoflagellate, and bacterial biomarkers were found in similar proportions in both zooplankton and particles with depth at both locations, providing additional evidence that mesopelagic zooplankton consume sinking particles. Carnivory indices were higher and increased significantly with depth at ALOHA, and exhibited distinct peaks at K2, representing an increase in dependence on other zooplankton for food in deep waters. Our results indicate that feeding ecology changes with depth as well as by location. These changes in zooplankton feeding ecology from the surface through the mesopelagic zone, and between contrasting environments, have important consequences for the quality and quantity of organic material available to deeper pelagic and benthic food webs, and for organic matter sequestration.     

Zooplankton vertical migration and the active transport of dissolved organic and inorganic carbon in the Sargasso Sea

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

Seston available as a food resource for the ribbed mussel (Geukensia demissa) in a North American,mid-Atlantic saltmarsh

We studied the composition of the <25 μm seston size fraction as a food resource potentially available to suspension feeding ribbed mussels, Geukensia demissa, over an annual cycle in Canary Creek saltmarsh, Delaware Bay. There were significant seasonal variations in the concentration of particulate organic carbon (POC), particulate organic nitrogen (PON), and total carbohydrate, but not cellulose. The concentration of cellulose, measured by hydrolytic cellulase enzyme assay, was relatively low (seasonal range 24 to 35 μg l⁻¹) and only comprised from 3% of total carbohydrate in May 1996 to 13% in November 1995. We used the biomass of microalgae, estimated from chlorophyll a, and abundance of free-living bacteria and heterotrophic nanoflagellates to calculate each component's equivalent carbon content. Microalgae were the most dominant carbon source (62% annually) among the four identified components (phytoplankton, bacteria, heterotrophic nanoflagellates, and cellulose) in all seasons except in August 1995 when carbon from bacteria was most abundant (55%). The annual average carbon equivalents of heterotrophic nanoflagellates and cellulose were relatively small (2 and 4%, respectively). The total concentration of POC in the seston was much greater than the carbon derived from the four identified components. The proportions that these identified components contributed to POC varied seasonally and combined only accounted for 8–24% of POC. Based on these estimates, the bulk of the POC in Canary Creek marsh was not associated with any of the four components we identified. We suggest that this uncharacterized material was some type of non-lignocellulosic, amorphous detritus of unknown utility as a food resource for ribbed mussels.     

Relationships between chlorophyll a, bacteria, ATP, POC and respiration rates in the Changjiang Estuary and the plume

Bacteria abundance, chlorophyll a, ATP and POC concentrations and respiration rates of microorganisms in the Changjiang Estuary and the plume were determined in July 1986. The high values of bacteria abundance occurred in the river mouth in association with suspended matter. It is assumed that bacteria were the major contributor to ATP and the main consumer of dissolved oxygen, and that the relationship between ATP and POC was present in that area. In the dilution zone (salinity; 25-30), instead of bacteria, phytoplankton was the major contributor to ATP and respiration rates, due to diatom bloom. Close relationships between Chi a and ATP, and ATP and POC were observed. Contribution of microbial carbon to POC was also estimated.     

Plankton distribution and vertical flux of biogenic matter during high summer stratification in the Krka estuary (Eastern Adriatic)

The aim of this study was to investigate phytoplankton abundance, composition and vertical export in the highly stratified Krka estuary, Croatia. The estuary is stratified throughout the year, and an interface between fresh- and brackish water plays an important role in production and degradation of biogenic matter. Vertical export of particulate organic carbon (POC), phytoplankton carbon (PPC) and faecal pellet carbon (FPC) was studied by deploying sediment traps in the middle and lower reach of the estuary and in the adjacent coastal zone. Zooplankton faecal pellet (FP) production experiments were conducted to provide additional information on the potential contribution of FP to the total carbon flux. High suspended concentrations of POC, chlorophyll a and phytoplankton was found in the lower reaches of the Krka estuary, adjacent to a source of anthropogenic eutrophication. The fraction of organic detritus to the total POC flux was 61–69% inside the estuary but only 7% at the marine station. This indicates that the primary producers in the surface layer of the Krka estuary are decomposed in and below the interface and then settle as detritus to the bottom. Low sedimentation rates in the coastal zone outside the estuary revealed that the eutrophication does not spread out of the estuary. Mesozooplankton played a modest role in vertical flux regulation, due to their low abundance and dominance of smaller forms as well as low faecal pellet production rates. It is concluded that processes taking place at the freshwater-seawater interface are of major importance for the vertical carbon flux in the investigated area.     

东海陆架典型断面颗粒态氨基酸的分布及控制因素分析

采用高效液相色谱法,通过现场调查对东海典型PN断面(文中为C断面)的颗粒态氨基酸(Particulate Amino Acids,PAA)进行了分析,并结合叶绿素a(Chla)、颗粒有机碳(POC)、颗粒有机氮(PON)及颗粒态氨基酸的构型特征(D和L型)等参数探讨了该区颗粒有机氮的来源和降解情况。结果表明,在长江口最大浑浊带,受咸淡水混合和生物现场生产双重作用影响,POC、PON以及PAA的总浓度均达到了极大值,其中,受再悬浮作用影响,底层水体中的有机物呈现高度降解的状态;近岸水华区域的颗粒态氨基酸则更多来源于现场生产,而且POC/Chla质量比值与降解因子DI值的负相关特征表明冲淡水向海洋输送的过程中,现场生产力对颗粒有机碳的贡献比重逐渐增大,悬浮颗粒物也变得越来越新鲜。值得关注的是,一些D型氨基酸[如D型天冬氨酸(D-Asp),D型丙氨酸(D-Ala)]与细菌生物量之间存在良好的正相关性,暗示颗粒态氨基酸在受到物理水团和生物现场生产作用控制的同时,还受控于微微型浮游生物以及异养细菌。     

南沙渚碧礁生态系有机碳的分布及周日变化特征

1999 年 4 月对我国南沙群岛渚碧礁海水中溶解有机碳的分布及礁坪区颗粒有机碳 (POC) 和溶解有机碳 (DOC) 的周日变化特征进行了观测。结果表明,渚碧礁表层海水 DOC 变化范围为 1.43~3.62 mg/L,平均为 2.16 mg/L,含量分布大致表现为礁坪区>潟湖>礁外。潟湖 DOC 的垂直分布大致表现为表层高于底层,可能与表层浮游植物的光合作用有关。礁坪区 POC 及 DOC 都呈现显著的周日变化特征,POC 呈现夜晚高,白天低的特点,浮游植物的昼夜垂直移动可能是产生该现象的主要原因。DOC 的周日变化则主要受浮游动物昼夜垂直移动及细菌等生物活动的影响。     

Food Preferences of Dominant Salt Marsh Herbivores and Detritivores

Abstract. Food preferences of six dominant salt marsh invertebrates were studied to identify detritivores and to assess differences in their diets. Animals fed on agar suspensions of dead and live foods in petri dishes with four compartments. Only two foods were included in each dish. Relative palatability was assessed by measuring the amount of suspension removed or by counting the number of feeding marks left on the surface of the suspensions.
Feeding marks reflected differences in mouth parts and feeding mechanisms of the six invertebrates. Melampus bidentatus, Orchestia grillus , and Philoscia vittata preferred dead litter over live tissues of marsh graminoids, blue-green algae, and sulfur bacteria. No preference for detritus from different graminoids was shown by M. bidentatus. Orchestia grillus fed preferentially on Spartina patens detritus. Philoscia vittata preferred detritus from S. alterniflora and S. patens. Blue-green algae and sulfur bacteria were preferred over detritus by Littorina saxatalis , but detritus was preferred over live graminoids. Littorina saxatalis fed preferentially on Juncus gerardi detritus. Blue-green algae, sulfur bacteria, and live graminoids were preferred over detritus by L. littorea. Talorchestia longicornis also preferred blue-green algae.
On the basis of their food preferences, Littorina saxatalis, Melampus bidentatus, Orchestia grillus , and Philoscia vittata were classified as detritivores. Feeding on detritus from different plant species could result in a partitioning of this food resource in the detritus-based food chains of the salt marsh ecosystem.     

Biogenic carbon flows through the planktonic food web of the Amundsen Gulf (Arctic Ocean): A synthesis of field measurements and inverse modeling analyses

Major pathways of biogenic carbon (C) flow are resolved for the planktonic food web of the flaw lead polynya system of the Amundsen Gulf (southeast Beaufort Sea, Arctic Ocean) in spring-summer 2008. This period was relevant to study the effect of climate change on Arctic marine ecosystems as it was characterized by unusually low ice cover and warm sea surface temperature. Our synthesis relied on a mass balance estimate of gross primary production (GPP) of 52.5 ± 12.5 g C m⁻² calculated using the drawdown of nitrate and dissolved inorganic C, and a seasonal f-ratio of 0.64. Based on chlorophyll a biomass, we estimated that GPP was dominated by phytoplankton (93.6%) over ice algae (6.4%) and by large cells (>5 μm, 67.6%) over small cells (<5 μm, 32.4%). Ancillary in situ data on bacterial production, zooplankton biomass and respiration, herbivory, bacterivory, vertical particle fluxes, pools of particulate and dissolved organic carbon (POC, DOC), net community production (NCP), as well as selected variables from the literature were used to evaluate the fate of size-fractionated GPP in the ecosystem. The structure and functioning of the planktonic food web was elucidated through inverse analysis using the mean GPP and the 95% confidence limits of every other field measurement as lower and upper constraints. The model computed a net primary production of 49.2 g C m⁻², which was directly channeled toward dominant calanoid copepods (i.e. Calanus hyperboreus 20%, Calanus glacialis 10%, and Metridia longa 10%), other mesozooplankton (12%), microzooplankton (14%), detrital POC (18%), and DOC (16%). Bacteria required 29.9 g C m⁻², a demand met entirely by the DOC derived from local biological activities. The ultimate C outflow comprised respiration fluxes (82% of the initial GPP), a small sedimentation (3%), and a modest residual C flow (15%) resulting from NCP, dilution and accumulation. The sinking C flux at the model limit depth (395 m) supplied 60% of the estimated benthic C demand (2.8 g C m⁻²), suggesting that the benthos relied partly on other C sources within the bottom boundary layer to fuel its activity. In summary, our results illustrate that the ongoing decline in Arctic sea ice promotes the growth of pelagic communities in the Amundsen Gulf, which benefited from a ∼80% increase in GPP in spring-summer 2008 when compared to 2004 – a year of average ice conditions and relatively low GPP. However, 53% of the secondary production was generated within the microbial food web, the net ecological efficiency of zooplankton populations was not particularly high (13.4%), and the quantity of biogenic C available for trophic export remained low (6.6 g C m⁻²). Hence it is unlikely that the increase in lower food web productivity, such as the one observed in our study, could support new harvestable fishery resources in the offshore Beaufort Sea domain.     

Carbon to nitrogen (C:N) stoichiometry of the spring–summer phytoplankton bloom in the North Water Polynya (NOW)

The carbon to nitrogen (C:N) stoichiometry of phytoplankton production varied significantly during the spring–summer bloom in the North Water Polynya (NOW), from April through July 1998. The molar ratio of particulate organic carbon (POC) to nitrogen (PON) production by phytoplankton (ΔPOC:ΔPON) increased from 5.8 during April through early June to 8.9 in late June and July. The molar dissolved inorganic carbon (DIC) to nitrate+nitrite (NO₃) drawdown ratio (ΔDIC: ΔNO₃) increased from 6.7 in April and May, to 11.9 in June (no estimate for July because of ice melting). The discrepancy between ΔPOC:ΔPON and ΔDIC:ΔNO₃ was likely due to dissolved organic carbon (DOC) production. Increased ΔPOC:ΔPON of phytoplankton and surface water ΔDIC:ΔNO₃ throughout the phytoplankton blooms resulted from changes in physical properties of the upper water column, such as reduced thickness of the surface mixed layer that exposed phytoplankton to increased photosynthetically available radiation (PAR), accompanied by NO₃ depletion. This is expected to have significant effects on the cycling of carbon (C) and nitrogen (N) in pelagic ecosystems, as the increased C:N ratio of organic matter decreases its quality as substrate for grazers and microbial communities. Based on ΔPOC:ΔPON, the ratio of POC to chlorophyll a (Chl) production (ΔPOC:ΔChl) and the relationship between Chl yields and NO₃ depletion, we estimate that 71±17% and 46±20% of the depleted NO₃ went to PON production in the euphotic zone over the polynya from April to early June, and late June to July, respectively. The remaining NO₃ was likely channelled to dissolved organic nitrogen (DON) and heterotrophic bacteria, which were not returned to the dissolved inorganic nitrogen (DIN) pool through recycling during the course of the study. Hence, the autotrophic production of organic N and its recycling by the microbial food web were not coupled temporally.     

Microbial respiration in the aphotic zone of the Ross Sea (Antarctica)

In January–February 2001, we measured microbial biomass as ATP and community respiration as ETS activity of organisms < 200 μm in the aphotic zone of the Ross Sea. Microbial respiration amounted to 2.14 mmol C m^− 2 day^− 1 in the depth range 200–1000 m. Our daily estimates of carbon export are close to the daily percentage of net community production (NCP), removed as sinking biogenic particles from the upper 100 m in the entire Ross Sea, but lower than those of other oceanic systems. Comparing remineralization determined in this study with that obtained by sediment traps in the Ross Sea, it appeared that about 63% of organic carbon remineralized by respiration derived from POC pool. Such evidence highlighted POC source as the main organic fuel of the biological pump in the Ross Sea.     

A new method of evaluating the mineralization of particulate and dissolved photoassimilated organic matter

A new method of evaluating the rate of mineralization of photoassimilated organic matter is described. This method enables us to compare the rate of direct mineralization of particulate organic carbon (POC) to CO₂ with the rate of solubilization of photoassimilated organic carbon followed by the mineralization of the resultant dissolved organic carbon (DOC) under the same conditions. The direct mineralization of photoassimilated carbon from POC to CO₂ is a more significant process compared with the mineralization of extracellular released organic carbon. The first-order rate coefficients range from 0.132 to 0.434 day^–1 for direct mineralization and 0.034 to 0.189 day^–1 for solubilization.     

Seasonal growth and senescence in continental shelf ecosystems: a test of the SEEP hypothesis

The hypothesis that continental shelf ecosystems export a major fraction of the carbon produced by the phytoplankton during the spring bloom was tested during the Shelf Edge Exchange Processes (SEEP) experiment off the north-east coast of the United States in 1984. The total benthic standing stocks in terms of organic carbon (macrofauna, meiofauna and bacteria) have been estimated in the SEEP area. Their preponderance on the continental shelf was partial evidence that little organic matter escapes to the upper continental slope. Measurements of the metabolism of the biota allowed calculation of turnover times of organic detritus and the total biota. The turnover time of detritus increased as grain size decreased, suggesting that fine-grained deposits contain mostly refractory, non-reactive compounds, especially on the deep slope. Turnover times of the total biota were about the same in the coarse- as in the fine-grained shelf deposits, but a far larger fraction of the turnover was attributed to the bacteria in the fine sediments. On average, about 25 per cent of the primary production appeared to be utilized by the aerobic benthos on the continental shelf in the SEEP area. The role of anaerobes at depth in the sediments remains uncertain. This study, along with a review of other recent work, leads to the conclusion that only a small fraction of continental shelf phytodetritus is exported across a distinct shelf-slope hydrographic frontal system. What is not consumed in the spring is utilized on the shelf during the ensuing stratified season. More open ended ecosystems may export production more readily.     

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

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

Carbon Isotope Geochemistry of the Orinoco Basin

Stable carbon isotope ratios have been used to study the sources of particulate organic carbon (POC) and total dissolved inorganic carbon in the Orinoco Basin. The isotopic composition of total dissolved inorganic carbon shows a range of from -8·1 to -23·0 ppt, an indication of dominance of biological processes. The isotopic composition of POC exhibits a range of from -24·1 to -34·6 ppt with little seasonal variation. The isotopic evidence indicates that the POC is predominantly of terrestrial origin rather than a result of in situ planktonic production. The similarity of isotopic composition of POC and coastal sediments suggests that riverine organic detritus has been transported 30-50 km offshore in a direction parallel to the Orinoco river channel.