Mechanisms leading to the frequent occurrences of hypoxia and a preliminary analysis of the associated acidification off the Changjiang estuary in summer

Based on literature data and shipboard observations, this study investigated the main environmental characteristics of the seafloor topography, current field, front, and upwelling that are closely related to hypoxia occurrence off the Changjiang estuary. The physical processes of the plume front and upwelling off the Changjiang estuary in summer were coupled. The vertical distribution pattern of the plume front was closely related to the upwelling. By reviewing and analyzing the historical summer hypoxia events off the Changjiang estuary, we statistically demonstrated the spatial structure of the horizontal distribution of the hypoxic zone and investigated the location of occurrence zone of the hypoxia. We found that the dissolved oxygen (DO) concentration on the inner continental shelf off the estuary showed a “V” shape in relation to station depth. Therefore, we noted that the hypoxic water on the inner continental shelf mostly occurred on the slopes with steep seafloor topography. Base on the current understanding of the hypoxic mechanisms off the Changjiang estuary, we analyzed the biogeochemical mechanisms that could cause the steep terrain off the Changjiang estuary to become the main areas prone to summer hypoxia and explained the internal relations between the location of the hypoxic zone on the slopes and the plume front and upwelling. The plume front and upwelling off the Changjiang estuary and their coupling were important driving forces of summer hypoxia. The continuous supply of nutrients affected by the interaction of the plume front extension of the Changjiang Diluted Water (CDW) and upwelling and the favorable light conditions were important mechanisms causing the phytoplankton blooms and benthic hypoxia off the Changjiang estuary in summer. By analyzing oxygen utilization, organic carbon mineralization, and nutrient regeneration in the hypoxic zone, we observed that the significant oxygen utilization process off the Changjiang estuary in summer also mainly occurred near the steep slopes with front and upwelling features and confirmed the apparent nutrient loss in the benthic hypoxic zone. Meanwhile, our analysis revealed that the sediment resuspension in the benthic boundary layer in the mud areas off the Changjiang estuary could also affect the oxygen utilization and mineralization of organic carbon and nutrient recycling and regeneration. This study also demonstrated that the steep terrain off the Changjiang estuary was the main location for summer acidification, and the coupling between the plume front and upwelling on the steep slopes was an important physical driving force inducing summer benthic acidification. Finally, we discussed issues to address in future studies of the hypoxic zone and water acidification off the Changjiang estuary.     

Oxygen depletion off the Changjiang (Yangtze River) Estuary

In a survey on the Yellow Sea and the East China Sea on August 20-30 of 1999, we found a hypoxic zone (<2 mg/L) of 13700 km² with an average thickness of 20m at the bottom of the Changjiang (Yangtze River) Estuary, with an oxygen minimum value of 1 mg/L. The extension of the dissolved oxygen deficiency extended to the 100m isobath in a southeastward direction along the bottom of the continental shelf of the East China Sea. During the last two decades, the minimum dissolved oxygen values in the low oxygen region of the Changjiang Estuary have decreased from 2.85 mg/L to 1 mg/L. In the hypoxic zone, the apparent oxygen utilization (AOU) was 5.8 mg/L and the total oxygen depletion approximately 1.59 × 10⁶t. The strong halocline above the hypoxic zone, as a result of affluent water from the Changjiang, Taiwan Warm Current (TWC), and the high concentrations of particle organic carbon (POC) and nitrogen (PON) are the major factors causing the formation of the hypoxic zone. The POC: PON ratios and nutrient concentration distributions in the hypoxic zone suggest that the oxygen deficiency in the bottom water during the summer in the East China Sea off the Changjiang is the result of organic carbon production enhanced by nutrients from the Changjiang and fluvial organic matter input, followed by a shift in regeneration of nutrients in the East China Sea.     

Comparison of hypoxia among four river-dominated ocean margins: The Changjiang (Yangtze), Mississippi,Pearl, and Rhône rivers

We examined the occurrence of seasonal hypoxia (O₂<2 mg l⁻¹) in the bottom waters of four river-dominated ocean margins (off the Changjiang, Mississippi, Pearl and Rhône Rivers) and compared the processes leading to the depletion of oxygen. Consumption of oxygen in bottom waters is linked to biological oxygen demand fueled by organic matter from primary production in the nutrient-rich river plume and perhaps terrigenous inputs. Hypoxia occurs when this consumption exceeds replenishment by diffusion, turbulent mixing or lateral advection of oxygenated water. The margins off the Mississippi and Changjiang are affected the most by summer hypoxia, while the margins off the Rhône and the Pearl rivers systems are less affected, although nutrient concentrations in the river water are very similar in the four systems. Spring and summer primary production is high overall for the shelves adjacent to the Mississippi, Changjiang and Pearl (1–10 g C m⁻² d⁻¹), and lower off the Rhône River (<1 g C m⁻² d⁻¹), which could be one of the reasons of the absence of hypoxia on the Rhône shelf. The residence time of the bottom water is also related to the occurrence of hypoxia, with the Mississippi margin showing a long residence time and frequent occurrences of hypoxia during summer over very large spatial scales, whereas the East China Sea (ECS)/Changjiang displays hypoxia less regularly due to a shorter residence time of the bottom water. Physical stratification plays an important role with both the Changjiang and Mississippi shelf showing strong thermohaline stratification during summer over extended periods of time, whereas summer stratification is less prominent for the Pearl and Rhône partly due to the wind effect on mixing. The shape of the shelf is the last important factor since hypoxia occurs at intermediate depths (between 5 and 50 m) on broad shelves (Gulf of Mexico and ECS). Shallow estuaries with low residence time such as the Pearl River estuary during the summer wet season when mixing and flushing are dominant features, or deeper shelves, such as the Gulf of Lion off the Rhône show little or no hypoxia.     

Seasonal changes of organic matter origins and anammox activity in the Changjiang Estuary deduced from multi-biomarkers in suspended particulates

Human activity-induced eutrophication and harmful algal blooms are main causes of the expansion of the hypoxic zone in the Changjiang Estuary. Among the many changes in biogeochemical processes, anaerobic ammonium oxidation (anammox) is proposed to play an important role in the nitrogen cycle in hypoxic areas. Ladderane lipids have been used as biomarkers to indicate anammox activity in ecosystems, but the origins of anammox bacteria and ladderanes in suspended particulates are still unclear. In this study, we report the results of a suite of biomarker analyses of suspended particulates across a salinity gradient of the Changjiang Estuary in both the spring and summer to evaluate the origins of the ladderanes and their potential as proxies for anammox activity and hypoxia. The spatio-temporal variations in terrestrial biomarkers (n-alkanes and n-alkanols), marine biomarkers (brassicasterol and dinosterol), and the Terrestrial and Marine Biomarker Ratio and Branched and Isoprenoid Tetraether indices reveal that marine organic matter was dominant in the particulates in both the spring (55%) and summer (86%) seasons. Correlations with both marine and terrestrial biomarkers suggest that ladderanes were mainly produced in the water column, and therefore that ladderane concentrations in suspended particulates in the Changjiang Estuary mainly reflect anammox activity in the water column, although changes in anammox bacterial assemblages may also have played a role in ladderane concentrations. Overall, ladderane results suggest that anammox activity was widespread in the Changjiang Estuary; but higher ladderane concentrations in the summer (especially in the upwelling zone) were correlated with lower dissolved oxygen concentrations, which suggest that they are useful proxies for hypoxia.     

Controls on the interannual variability of hypoxia in a subtropical embayment and its adjacent waters in the Guangdong coastal upwelling system,northern South China Sea

Coastal embayments located downwind of large rivers under an upwelling-favorable wind are prone to develop low-oxygen or hypoxic conditions in their bottom water. One such embayment is Mirs Bay, off the Guangdong coast, which is affected by upwelling and the Pearl River Estuary (PRE) plume during summer. The relative importance of physical and biochemical processes on the interannual variability of hypoxia in Mirs Bay and its adjacent waters was investigated using statistical analyses of monthly hydrographic and water quality monitoring data from 2001 to 2015. The results reveal that the southwesterly wind duration and the PRE river discharge together explain 49% of the interannual variability in the size of the hypoxic area, whereas inclusion of the nutrient concentrations inside Mirs Bay and phytoplankton on the shelf explains 75% of the interannual variability in the size of the hypoxic area. This finding suggests that the interannual variability of hypoxia in Mirs Bay is regulated by coupled physical and biochemical processes. Increase of the hypoxic area under a longer-lasting southwesterly wind is caused by increased stratification, extended bottom water residence time, and onshore transport of a low-oxygen water mass induced by stable upwelling. In contrast, a reduction in the size of the hypoxic area may be attributed to a decrease in the surface water residence time of the particulate organic matter outside Mirs Bay due to increased discharge from the PRE. The results also show that the effects of allochthonous particulate organic matter outside Mirs Bay on bottom hypoxia cannot be neglected.     

IndoTROPICS studies on the plume of the Mamberamo river into the Bismarck Sea, West Papua, Indonesia

The tropical river ocean processes in coastal settings (TROPICS) program in Indonesia (Indotropics) was carried out in the Mamberamo estuary on May–June 1999 and August 2000. The Mamberamo River flows northward from the high mountains of West Papua to the narrow and steep continental slope of the Pacific Ocean. The data for the 1999 Mamberamo estuary cruises show the dispersal of fresh water coming out of the Mamberamo River into the Bismarck Sea, and the plume is clearly defined by the pattern of salinity, turbidity, and nutrient distribution. The Mamberamo River and other nearby rivers supply high concentrations of phosphate and nitrate to the New Guinea Coastal Current, so that nutrient concentrations are higher in the surface estuarine plume, compared to offshore areas. The distribution of fishes and benthic animals were also sampled over this area.     

Distribution of chlorophyll-a off the Changjiang River and its dynamic cause interpretation

Red tide,also called harmful algal bloom interna-tionally,is a disaster abnormal phenomenon of oceanecology with an explosive breed or dense assemble ofone or several phytoplanktons in a specific ocean en-vironment condition,colors the seawater,influencesand harms ocean living things.The formation of redtide is controlled mainly by a complex interplay ofbiological,physical and chemical processes,but themost main cause influencing the occurrence of red tideis the seawater eutrophication,i.e.,the…     

Tidally induced upwelling off Yangtze River estuary and in Zhejiang coastal waters in summer

MASNUM wave-tide-circulation coupled numerical model (MASNUM coupled model, hereinafter) is de-veloped based on the Princeton Ocean Model (POM). Both POM and MASNUM coupled model are ap-plied in the numerical simulation of the upwelling off Yangtze River estuary and in Zhejiang coastal waters in summer. The upwelling mechanisms are analyzed from the viewpoint of tide, and a new mechanism is proposed. The study suggests that the tidally inducing mechanism of the upwelling in-cludes two dynamic aspects: the barotropic and the baroclinic process. On the one hand, the residual currents induced by barotropic tides converge near the seabed, and upwelling is generated to maintain mass conservation. The climbing of the residual currents along the sea bottom slope also contributes to the upwelling. On the other hand, tidal mixing plays a very important role in inducing the upwelling in the baroclinic sea circumstances. Strong tidal mixing leads to conspicuous front in the coastal waters. The considerable horizontal density gradient across the front elicits a secondary circulation clinging to the tidal front, and the upwelling branch appears near the frontal zone. Numerical experiments are de-signed to determine the importance of tide in inducing the upwelling. The results indicate that tide is a key and dominant inducement of the upwelling. Experiments also show that coupling calculation of the four main tidal constituents (M2, S2, K1, and O1), rather than dealing with the single M2 constituent, im-proves the modeling precision of the barotropic tide-induced upwelling.     

Effects of hypoxia on benthic organisms in Tokyo Bay, Japan: a review

Bottom hypoxia (dissolved oxygen concentration ≤2 ml l(-1)) from anthropogenic eutrophication is a growing global concern. Here, we summarized characteristics of hypoxia and its effects on benthic organisms in Tokyo Bay. Despite recent decreases in nutrient inputs, hypoxia has been increasing in duration and spatial extent, suggesting that the substantial loss of tidal flats from reclamation is contributing to a decrease in the ability of Tokyo Bay to recycle nutrients. Hypoxia develops in the central to northern part of the bay and persists from spring to autumn, causing defaunation of benthic organisms. After the abatement of hypoxia in autumn, the defaunated area is recolonized, either through migration or larval settlement. Some megabenthic species with a spawning peak in spring and summer experience failure of larval settlement, which is probably due to hypoxia. The adverse effects of hypoxia are an impediment to recovery of benthic organisms in Tokyo Bay.     

Interaction of a river plume with coastal upwelling in the northeastern South China Sea

Observational and modeling studies were conducted to investigate the Pearl River plume and its interaction with the southwesterly driven upwelling circulation in the northern South China Sea during the summer. After exiting the Pearl River Estuary, the discharged freshwater generates a nearly stationary bulge of freshwater near the entrance of the estuary. Forced by the wind-driven coastal upwelling current, the freshwater in the outer part of the bulge flows downstream at the speed of the current and forms a widening and deepening buoyant plume over the shelf. The plume axis gradually shifts offshore of the current maximum as a result of currents induced by the contrasting density at the nose of plume and by the intensified Ekman drift in the plume. In this plume–current system, the fraction of the discharged freshwater volume accumulated in the bulge reaches a steady state and the volume of newly discharged freshwater is transported downstream by the upwelling current. Enhancement of stratification by the plume thins the surface frictional layer and enhances the cross-shelf circulation in the upper water column such that the surface Ekman current and compensating flow beneath the plume are amplified while the shoaling of the deeper dense water in the upwelling region changes minimally. The pressure gradient generated between the buoyant plume and ambient seawater accelerates the wind-driven current along the inshore edge of the plume but retards it along the offshore edge. Along the plume, downward momentum advection is strong near the highly nonlinear source region and a weaker upward momentum advection occurs in the far field over the shelf. Typically, the plume is shaped by the current over the shelf while the current itself is adjusting to a new dynamic balance invoked by the plume-induced changes of vertical viscosity and the horizontal pressure gradient. The spatial variation of this new balance leads to a coherent change in the cross-isobath transport in the upper water column during upwelling.     

Effects of hypoxia on benthic organisms in Tokyo Bay,Japan: A review

Bottom hypoxia (dissolved oxygen concentration ⩽2 ml l⁻¹) from anthropogenic eutrophication is a growing global concern. Here, we summarized characteristics of hypoxia and its effects on benthic organisms in Tokyo Bay. Despite recent decreases in nutrient inputs, hypoxia has been increasing in duration and spatial extent, suggesting that the substantial loss of tidal flats from reclamation is contributing to a decrease in the ability of Tokyo Bay to recycle nutrients. Hypoxia develops in the central to northern part of the bay and persists from spring to autumn, causing defaunation of benthic organisms. After the abatement of hypoxia in autumn, the defaunated area is recolonized, either through migration or larval settlement. Some megabenthic species with a spawning peak in spring and summer experience failure of larval settlement, which is probably due to hypoxia. The adverse effects of hypoxia are an impediment to recovery of benthic organisms in Tokyo Bay.     

The seasonal variability of the northern Benguela undercurrent and its relation to the oxygen budget on the shelf

The seasonal variations in the advection and mixing of water masses in the northern Benguela were studied in relation to the oxygen minimum zone over the Namibian shelf. The used data set consists of hydrographic and current measurements from an oceanographic mooring 20 nm off Walvis Bay, monthly CTD transects from the Namibian 23°S monitoring line and recent large-scale hydrographic surveys. The current time series showed an intermittent southward continuation of the Angola Current (AC) through the Angola–Benguela frontal zone (ABFZ) into the northern Benguela, commonly known as poleward undercurrent. In austral summer hypoxic, nutrient rich South Atlantic central water (SACW) from the Angola Gyre is transported into the northern Benguela, whereas during the winter season the oxygen rich Eastern SACW (ESACW) spreads northward. The water mass analysis reveals a mixing between both water masses in the northern Benguela between the ABFZ and the Lüderitz upwelling cell (27°S). The oxygen balance over the Namibian shelf depends to a high extent on the water mass composition of the upper central water layer, controlled by the large scale and local circulation. The deviation of the measured oxygen concentration from its mixing concentration, calculated with the source water mass properties, was used to quantify the oxygen consumption. A new local definition SACW was derived to exclude biogeochemical processes, taking place in the Angola Gyre. The oxygen deficit in the northern Benguela central water amounted to about 60–80 μmol l⁻¹ at the shelf edge and increased up to 150 μmol l⁻¹ on the shelf, due to local oxygen consumption. In the austral summer anoxic bottom waters are observed at the central Namibian shelf, which correlate to an SACW fraction >55%. Periods with high SACW fraction in the water mass composition were congruent with hydrogen sulphide events detected by remote sensing.     

Physical–biological coupling in the Pearl River Estuary

The Pearl River Estuary is a subtropical estuary and the second largest in China based on discharge volume from the Pearl River. Processes in the estuary vary spatially and temporally (wet vs dry season). In the dry season at the head of the estuary, hypoxic and nearly anoxic conditions occur and NH₄ reaches >600 μM, NO₃ is ∼300 μM and nitrite is ∼60 μM indicating that nitrification and denitrification may be important dry season processes in the region extending 40 km upstream of the Humen outlet. There are very few biological studies conducted in this upper section of the estuary in either the dry or wet seasons and hence there is a need for further research in this region of the river. In the wet season, the salinity wedge extends to the Hongqimen outlet and oxygen is low (35–80% saturation). Nitrate is ∼100 μM, silicate ∼140 μM; and phosphate is relatively low at ∼0.5 μM, yielding an N:P ratio up to ∼200:1 in summer. Nutrients decrease in the lower estuary and primary productivity may become potentially P-limited. Eutrophication is not as severe as one would expect from the nutrient inputs from the Pearl River and from Hong Kong's sewage discharge. This estuary shows a remarkable capacity to cope with excessive nutrients. Physical processes such as river discharge, tidal flushing, turbulent dispersion, wind-induced mixing, and estuarine circulation play an important role in controlling the production and accumulation of algal blooms and the potential occurrence of hypoxia. Superimposed on the physical processes of the estuary are the chemical and biological processes involved in the production of the bloom. For example, the 100N:1P ratio indicates that P potentially limits the amount of algal biomass (and potential biological oxygen demand) in summer. While extended periods of hypoxia are rare in Hong Kong waters, episodic events have been reported to occur during late summer due to factors such as low wind, high rainfall and river discharge which result in strong density stratification that significantly dampens vertical mixing processes. Nutrient loads are likely to change over the next several decades and monitoring programs are essential to detect the response of the ecosystem due to the future changes in nutrient loading and the ratio of nutrients.     

Impact of hypoxia on the structure and function of benthic epifauna in Tolo Harbour,Hong Kong

In Tolo Harbour, Hong Kong, where hypoxia occurs periodically, a large amount of benthic epifauna was found to be absent in the summer but become abundant again in the winter. This observation may indicate that some species are sensitive in detecting low dissolved oxygen (DO) levels and are able to escape from the harbour to more oxygenated, open waters when conditions deteriorate and return when DO levels are improved. Analysis of biological traits of these animals showed that 58% of the categories showed significant differences between the community structure in summer and winter. The significant biological traits of the few species, which remained in the harbour under the summer’s hypoxic conditions suggested adaptation rather than opportunism to the low DO levels. Our data also showed that trawling can increase DO at the bottom and may benefit the less sensitive species that remained in the habitat during the summer.     

Impact of hypoxia on the structure and function of benthic epifauna in Tolo Harbour, Hong Kong

In Tolo Harbour, Hong Kong, where hypoxia occurs periodically, a large amount of benthic epifauna was found to be absent in the summer but become abundant again in the winter. This observation may indicate that some species are sensitive in detecting low dissolved oxygen (DO) levels and are able to escape from the harbour to more oxygenated, open waters when conditions deteriorate and return when DO levels are improved. Analysis of biological traits of these animals showed that 58% of the categories showed significant differences between the community structure in summer and winter. The significant biological traits of the few species, which remained in the harbour under the summer's hypoxic conditions suggested adaptation rather than opportunism to the low DO levels. Our data also showed that trawling can increase DO at the bottom and may benefit the less sensitive species that remained in the habitat during the summer.     

Microbial food web contributions to bottom water hypoxia in the northern Gulf of Mexico

Nutrients from the Mississippi/Atchafalaya Rivers greatly stimulate biological production in the ‘classical’ food web on the inner shelf of the northern Gulf of Mexico. Portions of this production, especially large diatoms and zooplankton fecal pellets, sink and decompose in the bottom water, consuming oxygen and contributing to the annual development of an extensive zone of bottom water hypoxia, typically >15,000 km² since 1993. The microbial food web is also active in the Mississippi River plume, but consists of small organisms that sink slowly. This ‘recycling’ food web has not been considered as a significant contributor to vertical flux and hypoxia. However, gelatinous zooplankton, especially pelagic appendicularians such as Oikopleura dioica, mediate the conversion of microbial web organisms to organic particles with high sinking rates. When pelagic appendicularians are abundant in coastal regions of the northern Gulf of Mexico, they stimulate the rapid vertical transfer of microbial web productivity in the surface layer, which is only 5–15 m thick in the coastal hypoxic region, to the sub-pycnocline layer that becomes hypoxic each summer. In this paper we present results from two studies examining the significance of this pathway. In both 2002 and 2004, we observed high production rates of appendicularians in coastal waters. Discarded gelatinous houses and fecal pellets from the appendicularian populations often provided more than 1 g m⁻² d⁻¹ of organic carbon for the establishment and maintenance of hypoxia in the northern Gulf of Mexico. This source of organic matter flux is especially important in regions far from the river plumes and during periods of low river discharge. Autotrophic elements of this food web are primarily supported by recycled inorganic nutrients originating in the Mississippi and Atchafalaya Rivers. Sources of dissolved organic matter (DOM) supporting the heterotrophic components of this microbial food web may include in situ production, the Mississippi/Atchafalaya Rivers, and Louisiana's coastal wetlands. If significant, the latter source provides a possible link between Louisiana's high rates of coastal land loss and the large hypoxic zone observed along the coast during summer. Both of the latter DOM sources are independent of phytoplankton production stimulated by inputs of riverine inorganic nutrients.     

Primary production in a turbid estuary, the Ems-Dollard: field and model studies

The Ems-Dollard Estuary has a steep axial gradient in turbidity which strongly regulates primary production of phytoplankton and microphytobenthos. A combination of the turbidity and nutrient gradients causes a high pelagic primary production in the outer, seaward part of the estuary, and a high benthic production on high-level mudflats (with long effective photoperiods) in the inner part. Model simulations were used to study phytoplankton succession and the factors influencing this succession. Light limitation was evaluated in a sensitivity run which used a lower compensation light intensity.     

Nutrient processes and chlorophyll in the estuaries and plume of the Gulf of Papua

This paper investigates the waters of the Gulf of Papua during three cruises of the TROPICS (Tropical River Ocean Processes In Coastal Settings) programme. Plume characteristics were investigated during Leg 1 (May 1997), and estuarine properties during Leg 5a (September 1997) and Leg 7 (January 1999). During Leg 1 the plume was apparent as a well mixed layer up to 30 m deep extending offshore to a distance of 150 km off the Fly River. Lowest salinities were found off the Taruma Delta. Highest chlorophyll concentrations were found at the inner plume close to the river mouth. Dissolved phosphate and nitrate are removed in this zone, whereas silicate behaves conservatively. Primary productivity within the plume appears to rely upon recycled nutrients, with organic fractions representing the majority of the nutrient pool. In the estuaries nutrients were found to behave differently during the monsoon than during the low flow of the extremely dry conditions associated with the 1997 El Niño event. Normally the Fly is characterised by remineralisation of organic nitrogen in the upper estuary, but during drought conditions DON production and NH₄⁺ uptake suggest that bacterial activity is more prevalent. Ocean Colour and Temperature Scanner imagery shows a number of features of the plume, but generally overestimates chlorophyll concentrations due to the effects of high suspended sediment concentrations and, to a lesser extent, coloured dissolved organic matter.     

Modeling the response of primary production and sedimentation to variable nitrate loading in the Mississippi River plume

Increases in nitrate loading to the Mississippi River watershed during the last 50 years are considered responsible for the increase in hypoxic zone size in Louisiana-Texas shelf bottom waters. There is currently a national mandate to decrease the size of the hypoxic zone to 5000 km² by 2015, mostly by a 30% reduction in annual nitrogen discharge into the Gulf of Mexico. We developed an ecosystem model for the Mississippi River plume to investigate the response of organic matter production and sedimentation to variable nitrate loading. The nitrogen-based model consisted of nine compartments (nitrate, ammonium, labile dissolved organic nitrogen, bacteria, small phytoplankton, diatoms, micro- and mesozooplankton, and detritus), and was developed for the spring season, when sedimentation of organic matter from plume surface waters is considered important in the development of shelf hypoxia. The model was forced by physical parameters specified along the river-ocean salinity gradient, including residence time, light attenuation by dissolved and particulate matter, mixed layer depth, and dilution. The model was developed using measurements of biological biomasses and nutrient concentrations across the salinity gradient, and model validation was performed with an independent dataset of primary production measurements for different riverine NO₃ loads. Based on simulations over the range of observed springtime NO₃ loads, small phytoplankton contributed on average 80% to primary production for intermediate to high salinities (>15), and the main contributors to modeled sedimentation at these salinities were diatom sinking, microzooplankton egestion, and small phytoplankton mortality. We investigated the impact of limiting factors on the relationship between NO₃ loading and ecosystem rates. Model results showed that primary production was primarily limited by physical dilution of NO₃, followed by abiotic light attenuation, light attenuation due to mixing, and diatom sinking. Sedimentation was mainly limited by the first three of these factors. Neither zooplankton grazing or plume residence times acted as limiting factors of ecosystem rates. Regarding nutrient reductions to the watershed, simulations showed that about half of the percent decrease in NO₃ load was reflected in decreased plume sedimentation. For example, a 30% decrease in NO₃ load resulted in a 19% decrease in average plume primary production and a 14% decrease in sedimentation. Finally, our model results indicated that the fraction of primary production exported from surface waters is highly variable with salinity (7–87%), a finding which has important implications for predictive models of hypoxic zone size that assume a constant value for this ratio.     

Long-term photographic,current, and nephelometer observations of manganese nodule environments in the Pacific

Long-term measurements on and near the seafloor were made at five locations in the Pacific as part of the Manganese Nodule Project (MANOP). Bottom Ocean Monitor (BOM) tripods deployed at each site contained a time-lapse camera, nephelometer, and current meter. From nearly three years of photographs of the seafloor, there was no evidence that benthic organisms turned over, rotated, or moved nodules. There were indications, however, that benthic organims cleaned sediment from the tops of nodules and in some cases deposited sediment underneath nodules. Currents never exceeded the expected threshold for sediment resuspension of nodule movement. Particle concentrations in the water remained uniformly low throughout the deployments. Degredation of fecal pellets and organic matter proceeded more rapidly than expected. All changes of the seafloor morphology were attributable to the activity of organisms as opposed to currents. The occurrence in one photo sequence of dark, fluffy globs may be related to phytoplankton blooms in the region of equatorial upwelling.