Flows of materials between poorly flooded tidal marshes and an estuary

Flows of particulate carbon, nitrogen, phosphorus, chlorophyll a, crude fiber, carbohydrate, and adenosine tri-phosphate; and of dissolved nitrogen and phosphorus between a marsh and the Patuxent estuary, Maryland, USA, were measured over a 2-year period. Virtually no carbon was exchanged, while net flows of nitrogen and phosphorus were from the marsh to the estuary, principally in dissolved forms.Contribution No. 664, Center for Environmental and Estuarine Studies, University of Maryland.     

Energy flow through the benthos in a Scottish Sea Loch

Analysis of material collected in settlement traps at regular intervals over a period of a year gave an estimate of the annual organic input to the bottom sediment of about 28 gC/m² year, which is a little less than one third of the primary production in the overlying water column. The aerobic benthic community metabolism, estimated from in situ respiration measurements, was not significantly different from the carbon input. The rate of release of ammonia from the sediment was also measured in situ and would be sufficient to supply the greater part of the required input of inorganic nitrogen for photosynthesis in the water column.     

Carbon budget for the spring bloom in Auke Bay,Alaska

This paper describes a carbon budget for the spring phytoplankton bloom in Auke Bay, a subarctic bay in southeastern Alaska. The budget was constructed using semiweekly data on carbon production, particulate carbon in the water column, and cumulative sedimentation of carbon, chlorophyll a, and pheopigments. From these measured parameters, seasonal carbon consumption, utilization, and import/export terms were derived. The chlorophyll and pheopigment data were used to partition carbon sinking out of the photic zone between phytoplankton cells and fecal material. The difference between total carbon production and carbon available for consumption was attributed primarily to carbon import/export related to advection of water masses into and out of the bay. Separate budgets were developed for each of five sampling years (1985–1989). An average of 130±16 g C/m² were produced by phytoplankton during each spring. Our model suggests that an average of 70% of this carbon was available for consumption by grazers within the bay; the remaining 30% is assumed to have been exported from the bay by advective transport. Of the available (non-exported) carbon, an average of 55% was consumed by grazers, 34% sank out of the photic zone in the form of uneaten algae, and about 11% remained at the end of the sampling period in the form of phytoplankton standing stocks. Overall, about 27% of the carbon produced each spring in Auke Bay (35 gC/m²) was used for growth and respiration by first-order consumers within the bay.     

Benthic oxygen uptake and carbon cycling under aphotic and resource-limiting conditions in a submarine cave

To establish relationships between organic input to the benthos and decreases in benthic population biomass and density, benthic oxygen uptake was measured in an oligotrophic submarine cave in the northwestern Mediterranean Sea (Marseille, France), on seven separate occasions in 1987, using an in situ bell-jar respirometer. Oxygen uptake was measured in both the outer twilight section and the dark inner section of the cave during an annual survey (seven recording periods from February 1987 to November 1988). The mean annual benthic oxygen uptake was 80.9 litres O₂ m^–2 yr^–1 for the twilight outer section and 15.5 litres O₂ m^–2 yr^–1 for the dark inner section. The results are discussed and the biogeochemical budget for particulate organic carbon at the sediment-water interface calculated. Respiration rates (expressed as carbon equivalents), together with previously published data on vertical fluxes and burial of organic carbon, revealed that anaerobic pathways accounted for 14% and aerobic pathways for 86% of the total benthic metabolism in the outer part of the cave. In the inner section of the cave, degradation of organic carbon occurred only through aerobic degradation, indicating a strongly carbon-limited ecosystem. The low respiration rates recorded in the dark section were similar to values recorded for some oligotrophic deep-sea environments (1 000 to 2 000 m). Such budgets are essential preliminary steps in order to accurately model benthic metabolic pathways. The determination of annual fluxes linked to the acquisition of long-term data will yield better knowledge of the recycling processes at the water-sediment interface.     

Benthic community respiration in the N.W. Atlantic Ocean: in situ measurements from 40 to 5200 m

Benthic community respiration was measured in situ at 9 stations along the Gay Head-Bermuda transect from depths of 40 to 5200 m. Three methods were used; bell jar respirometers, grab respirometers, and free vehicle respirometers. Benthic community respiration rates spanned three orders of magnitude, decreasing from 21.5 ml O₂ m^-2 h^-1 at 40 m in November to 0.02 ml O₂ m^-2 h^-1 at 5200 m. Rates decreased two orders of magnitude between 40 and 1800 m and then significantly declined again between the continental rise (3650 m) and the abyssal plain stations. Predictive equations for benthic community respiration along the transect reflect a strong correlation with depth of water. Of lesser significance are the correlations with water temperature, dissolved oxygen, benthic animal biomass, surface primary productivity and sediment organic matter. Calculations show that annual benthic respiration can utilize 1 to 2% of the surface primary productivity. Of the 2 to 7% organic carbon fixed at the surface which supposedly reaches the bottom, only 15 to 29% is utilized by the benthic community at 2200, 3000, and 3650 m. The energy requirements of other biological components of deep-sea benthic communities, such as benthopelagic and macro-epibenthic animals, not included in these measurements, must also be considered in calculating a balance of carbon.Contribution from Scripps Institution of Oceanography.     

Effects of graveling on the primary productivity,respiration and nutrient flux of two estuarine tidal flats

This study was conducted in Puget Sound, USA, and investigated the effect of graveling intertidal mud and sandflats to enhance clam production on the benthic assemblage structure, primary productivity, respiration, and nutrient flux. The study was conducted between spring and autumn (1991), the period of greatest productivity and plant standing stock in Puget Sound. Graveled and control plots were established in the low intertidal zone on a mud flat in a protected embayment (Chapman Cove) and an exposed sandflat (Semiahmoo Bay). Gravel altered benthic assemblage structure, respiration, and nutrient flux rates. Graveled plots contained more surface coverage of sessile animals and seaweeds. Net productivity (NP), which differed relatively little between graveled and control plots, was negative for all but one sampling at the protected embayment plots. In contrast, NP was always positive at the exposed sandflat plots. The respiration rate was 13 to 57% greater in the graveled plot at Chapman Cove than in the adjacent control plot, and 7 to 54% greater in the gravel plot than the control plots at Semiahmoo Bay. Heterotrophy was greater in the graveled plots, as reflected by a lower net productivity to respiration ratio. Effects of graveling on water quality parameters such as dissolved oxygen and inorganic nitrogen concentrations were not detected. Graveling sandy and mud beaches increases secondary productivity, which is associated with increased rates of remineralization and release of dissolved nutrients to the water column.     

Nutrient cycling between the water column and a marine sediment. I. Organic carbon

The carbon flow through the sediments at a station located in 18.3 m of water off the Scripps Institution of Oceanography, San Diego, California (USA) was determined. The parameters studied [and their mean rates of input (+) or output (-) to the benthos] were macro-detritus (+0.028 gC m^-2day^-1), fallout of particulate debris (+3.3 gC m^-2day^-1), benthic net photosynthesis during the day (-0.06 gC m^-2 daylight period^-1), burial (0 gC m^-2day^-1), benthic respiration at night (-0.28 gC m^-2 night period^-1), and resuspension (-3.0 gC m^-2day^-1). Resuspension of sediment at this station was found to have a controlling effect on the sediment organic carbon content. Benthic photosynthesis was able to provide 79% of the organic carbon required by the benthos for respiration during the daylight hours. A carbon-flow diagram linking together all of the above measurements is presented.     

In situ filtering and ingestion rates of deep-sea benthic boundary-layer zooplankton in the Santa Catalina Basin

In situ rates of filtration, particulate ingestion, and carbon ingestion of deep-sea benthic boundary-layer zooplankton were determined in December 1984 in the Santa Catalina Basin, at 1 300 m depth in the California Borderland, by a short-term radioisotope-incorporation technique. Zooplankton were collected at 1 or 50 m above the bottom with an opening-closing net system on a submersible, and incubated at depth with labelled amino acids in special cod-end chambers. Concentrations of particulate material and particulate organic carbon in the ambient water were also measured. The zooplankton had a median weight-specific filtration rate of 12.4 ml (mg dr. wt)^-1 h^-1 and a median carbon ingestion rate of 5.4 g C (mg dr. wt)^-1 h^-1. Filtration rates were not significantly different from those in similar experiments in the north Atlantic at 2 175 m depth or Narragansett Bay in the winter, although the medians of the deep-sea experiments were lower than for the Bay. In the Santa Catalina Basin, rates from experiments at 1 m above the bottom in more turbid water were not significantly different from those at 50 m above the bottom in clearer water. These deep-sea benthic boundary-layer zooplankton may have the potential to respond to food pulses, and their relatively high ingestion rates suggest that they could have significant effects on particulate, chemical, and bacterial processes in the near-bottom water column.     

Decomposition of urea associated with photosynthesis of phytoplankton in coastal waters

Decomposition of urea in seawater was studied in Mikawa Bay, a shallow eutrophic bay on the southern coast of central Japan. The urea concentration in seawater ranged from 1.3 to 5.9 μg-at. N/1 and comprised 12 to 40% of the dissolved organic nitrogen. Using ¹⁴C labelled urea, the rate of CO₂ liberation from urea and the incorporation rate of urea carbon into the particulate organic matter were determined. For the surface samples, high rates of CO₂ liberation from urea as well as the incorporation of urea carbon into the particulate organic matter were observed in the light, while much lower rates were obtained in the dark. Incubation experiments with exposure to different light intensities revealed that the rate of CO₂ liberation from urea and the incorporation of urea carbon into particulate organic matter changed with light intensity, showing a pattern similar to that of photosynthesis. The highest liberation and incorporation rates were observed at 12,000 lux. Incubation in light and in dark produced marked decreases and increases, respectively, in urea and ammonia, while no appreciable changes were observed for nitrate and nitrite. It is suggested that urea decomposition associated with photosynthetic activity of phytoplankton is one of the major processes of urea decomposition, and that it plays a significant role in the nitrogen supply for phytoplankton in coastal waters.     

Decomposition of faecal matter and somatic tissue of Mytilus galloprovincialis: changes in organic matter composition and microbial succession

Variations in the biochemical composition of pseudo-faeces and faeces egested by Mytilus galloprovincialis (Lamarck) and in detritus derived from the somatic tissue of mussels during the decomposition process were investigated by means of two intensive experiments. During the degradation process, the biochemical composition of pseudo-faeces and faeces showed a clear increase in protein content related to the microbial colonization. Changes also occurred in the biochemical composition of particulate organic matter (POM) in the surrounding water due to faecal matter decomposition, heterotrophic utilization and conversion of particulate carbohydrates and proteins to the dissolved pool. The study of production and heterotrophic utilization of the POM derived from the somatic tissue of M. galloprovincialis collected in the Gulf of Tigullio, Italy in 1990 indicates that this kind of material is rapidly decomposable and largely available for benthic consumers. Bacteria utilized selectively the different compounds, and proteins proved to be the most suitable substrate for bacterial growth. The input of organic detritus into the experimental system resulted in an enhancement of bacterial activity and consequently of the RNA/DNA ratio. Bacterial DNA accounted on average for 17% of particulate DNA. During decomposition processes, nutrient release was about ten times higher than the value calculated from individual excretion rates, indicating that mussel beds may be important sites for nutrient regeneration. Carbon conversion efficiency for bacteria growing on faecal matter was, on average, 17.2%. The potential importance of faecal output and bacterial production as a carbon resource for benthic communities near mussel culture areas is discussed.     

The significance of microbial carbon in the nutrition of the deposit feeding polychaete Nereis succinea

A partial carbon budget was calculated for a population of the deposit feeding polychaete Nereis succinea (Frey and Leuckart) for a North Carolina, USA salt marsh in order to determine if the ingestion and assimilation of microbial carbon was sufficient to meet the carbon requirement. Carbon required by the population was estimated by calculating annual production, 2.1 g C m^-2, and respiration, 9.4 g. There was no net release of dissolved organic carbon. Annual consumption of microbial carbon (as determined by ATP) was estimated to be 5.2 g m^-2. Assimilation efficiency of heterotrophic, detrital microbes was estimated to be 57%. If this value is used for all the microbial carbon, then total assimilation was 3.0 g C m^-2, or about one-fourth the carbon requirement. N. succinea was able to assimilate carbon from sterile plant detritus which suggests that some of the carbon needed to balance the budget may come from direct uptake of the plant substrate. Other possible additional sources of carbon include consumption of meiofauna and uptake of dissolved organic matter.     

Suspended particulate organic matter in a Mediterranean submarine cave

Submarine caves display a paucity of benthic density and biomass that may be related to low trophic resources. Analysis of organic carbon, organic nitrogen, carbohydrate, protein and lipid content of suspended particulate matter was made during the period July 1985–July 1987 in a Mediterranean cave (Marseille, France) in order to determine any differences in the particulate organic matter composition along an horizontal transect. Particulate organic matter content clearly declined from the entrance of the cave to the dark inner area. This impoverishment could not be explained by a simple decrease in a few organic compounds, but appeared to be related to the combination of a decrease in both the amount and the composition of the suspended particles. Three progressive levels of impoverishment were identified towards the dark inner area of the cave: (i) decreasing amounts of seston; (ii) decreasing organic content of particles; (iii) increasing proportions of the geopolymeric (i.e., humic) components in the remaining organic matter, indicating increased degradation. The cave appeared to be sharply divided into two distinct sections — a twilight outer section whose waters contained slightly lower amounts of particulate organic matter than the open sea, and a dark inner section, 8 to 10 m higher, separated from the outer section by a steep rise and with waters of low organic matter content. The water in the twilight section appeared to be in thermal equilibrium with the open sea, and that in the dark inner section displayed thermal stratification. These differences indicated the presence of two distinct water bodies with contrasting average residence times, estimated as 1 d in the outer twilight section and 8 d in the dark inner section. The joint action of sedimentation and degradation resulted in an abrupt depletion of particulate organic matter in the dark inner section accompanied by a decrease in the benthic fauna. The decline in benthic heterotroph populations is probably related to the abrupt transition to oligotrophic conditions.     

Behavioral response of fish larvae to low dissolved oxygen concentrations in a stratified water column

Density stratification and respiration lead to vertical gradients in dissolved oxygen in many aquatic habitats. The behavioral responses of fish larvae to low dissolved oxygen in a stratified water column were examined during 1990–1991 with the goal of understanding how vertical gradients in dissolved oxygen may directly affect the distribution and survival of fish larvae in Chesapeake Bay, USA. In addition, the effects of low oxygen on 24-h survival rates were tested so that results of behavior experiments could be interpreted in the context of risk to the larve. Naked goby [Gobiosoma bosc (Lacépède)] and bay anchovy [Anchoa mitchilli (Valenciennes)] larvae strongly avoided dissolved oxygen concentrations <1 mg 1^-1, which were lethal within 24 h at 25 to 27°C. In addition, naked goby larvae, whose behavior was tested at a wider range of dissolved oxygen concentrations, also showed a reduced preference for an oxygen concentration of 2 mg 1^-1, which leads to reduced survival during long-term exposures and to reduced feeding rates. There were no major differences in behavior or survival between the two species, or between the two age classes of naked gobies tested. Results suggest that behavioral responses to oxygen gradients will play a large role in producing marked vertical changes in abundance of feeding-stage larvae in Chesapeake Bay; mortality from direct exposure to low oxygen will likely be much less important in producing vertical patterns of larval abundance.     

Particulate organic carbon and nitrogen in the eastern pacific ocean

Particulate organic carbon and nitrogen in sea water were measured in samples collected along a line 155°W; 50° N-15°S, during the cruise of R.V. “Hakuho-Maru” (KH-69-4). High concentrations of particulate matter were generally found at or near the sea surface; the concentrations decreased rapidly with depth. A consistent minimum was located in the depth range 150 to 250 m through the entire section sampled. The subsurface maximum layers roughly coincided with the chlorophyll maximum, but several irrregularities were noted. One of the most remarkable features of the vast stratum below 200 m depth was the presence of distinctively regional variation in concentration of particulate material through the entire section. In the section, we could define at least 6 large water parcels, vertically oriented, all with significantly different concentrations of both carbon and nitrogen. Variation in the deep water ranged from less than 5 μgC/l to more than 50 μgC/l. Correlation analysis between carbon concentration and apparent oxygen utilization (AOU) of ambient water for all samples showed that the carbon from particle-poor water parcels consistently decreased with increasing AOU, levelling to a practically constant low of around 5 to 10 μgC/l, whereas the carbon from particle-rich parcels was anomalously high (10 to 50 μgC/l) in the range of high AOU, and showed no consistent trend of convergence. The intergrated amount of particulate carbon in the total water column at each station was in the range 20 to 150 gC/m². More than 90% of this total amount was in the water column below 200 m depth, and the correlation of total amount of particulate material between the surface layer (0 to 200 m) and the water column below 200 m depth was highly significant. These observations are considered to indicate that the downward transport of these materials may be much quicker than so far estimated, at least in some localized areas.     

Ingestion of natural particulate organic matter and subsequent assimilation,respiration and growth by tropical lagoon zooplankton

Rates of ingestion of natural particulate organic matter and subsequent assimilation and respiration by zooplankton at Enewetak Atoll lagoon (Marshall Islands) were measured using a flow-through system. Maximum daily ingestion rates of carbon and nitrogen, expressed as a percentage of the body content, were 79 and 37%, respectively, for the large copepod Undinula vulgaris; 112 and 65%, respectively, for a group of mixed small copepods; and 61 and 34%, respectively, for the pteropod Creseis acicula. Daily metabolic carbon losses, expressed as above, were 63% for U. vulgaris, 88% for the small copepods, and 50% for C. acicula. Assimilation efficiences of carbon and nitrogen ranged from about 86 to 91%. The above rates are generally higher than in previous reports for similar sized zooplankton in temperate waters, while the daily growth increments, expressed as a percentage of the body carbon content (4.8% for U. vulgaris, 8.6% for the small copepods, and 2.6% for C. acicula), are comparable. It appears that the high rates of ingestion and assimilation of organic matter are compensated by high metabolic losses. These results indicate that at least for carbon, tropical zooplankton may have low growth efficiencies ranging from 4 to 9%.     

Feeding strategies of some demersal fishes of the continental slope and rise off the Mid-Atlantic Coast of the USA

Stomach contents of 729 fishes comprising 16 species were examined from the continental slope and rise off the Middle Atlantic States of the USA. Two main feeding modes among demersal deep-sea fishes were evident: those feeding primarily on pelagic food items, and those feeding on benthic invertebrates. Both pelagic and benthic predators were euryphagous. Most pelagic predators also fed on the epibenthos. These findings support Dayton and Hessler's (1972) suggestion that benthic predators should have a generalized diet which may be responsible for the high diversity found in the deep-sea infauna. The mesopelagic fauna is an important food source for some demersal fishes on the continental slope. Pelagic prey, which are also important to ecologically dominant demersal fishes on the lower slope and continental rise, may be nutritionally supported by suspended particulate organic matter in a nepheloid layer close to the bottom, and they may exist in much higher concentrations than in the bathypelagic zone above.Virginia Institute of Marine Science Contribution No. 835.     

In situ grazing rates of deep-sea benthic boundary-layer zooplankton

In situ grazing rates of mixed assemblages of deep-sea benthic boundary-layer zooplankton were measured in July 1983 at a site in the North Atlantic Ocean at 2 175 m depth using a short-term radioisotope-uptake method. Zooplankton were collected with an opening-closing net system from the bottom 1 m of the ocean and incubated in situ with mixed tritiated amino acids in special cod-end chambers. Incubations were terminated at depth by the addition of MS-222. Radioisotope uptake beyond that of dead controls was shown by both the zooplankton and particulate fractions. Grazing rates in the deep-sea experiments were surprisingly high, being comparable to wintertime Narragansett Bay zooplankton grazing rates determined in a separate series of laboratory experiments. These laboratory experiments also documented nonparticle-associated uptake of dissolved amino acids by Narragansett Bay zooplankton, but the importance of this in the deep sea is unknown. The deep-sea benthic boundary layer may be a region of elevated rates and activity because of its higher particulate concentrations, and our experiments may also have measured maximum rather than average rates.     

A model of macroinvertebrate trophic structure and oxygen demand in freshwater wetlands

We present a model of macroinvertebrate trophic structure, detrital cycling, and dissolved oxygen (DO) dynamics in shallow freshwater wetlands with varying allochthonous subsidies. The model is based on field data of primary and secondary production in municipal wastewater-fed and river-fed constructed wetlands in central Ohio, USA. State variables for primary production include macrophyte, periphyton, and metaphyton biomass. Macroinvertebrate biomass is segregated by functional feeding group and includes collectors, scrapers, shredders, and predators. Model simulations demonstrate the association of water column dissolved oxygen, primary production, allochthonous organic matter, and the structure of the macroinvertebrate community. The quality and quantity of allochthonous carbon is shown to have considerable importance, not only as a food source but also as an oxygen sink. Allochthonous carbon equivalent to 5% of autochthonous production increases the macroinvertebrate standing crop by 4–17%, depending on particle size. A large allochthonous subsidy also reduces the simulated average diel dissolved oxygen and increases the percentage of hypoxia-tolerant macroinvertebrates. Simulations show both the heterotrophic response and the changes in community structure brought about by an allochthonous subsidy.     

Sedimentation and benthic mineralization of organic detritus in a Norwegian fjord

The supply of particulate material to the sea-bed as well as the oxygen consumption and the redox potential of the sea-bed were measured during a one-year period (1979/1980) at 60 and 90 m depth in the inner part of a west Norwegian fjord, Fanafjorden. At both sites, uniform sedimentation rates of total particulate material (825 and 885 g m^-2 yr^-1, respectively) and particulate inorganic material (576 and 616 g m^-2 yr^-1, respectively) were found. The sedimentation rates of particulate organic carbon (96 and 107 g m^-2 yr^-1, respectively) and particulate organic nitrogen (10 and 12 g m^-2 yr^-1, respectively) were low in winter, higher in summer and autumn, with maxima in May/June, reflecting similar maxima in the phytoplankton biomass in the area, with 6 to 8 wk delay. The oxygen consumption of the sea-floor was lowest in winter/spring and highest in summer. Thirtytwo and 38 g C m^-2 yr^-1 (respiration quotient=0.85) were metabolized by the sediment at 60 and 90 m, respectively. The simultaneous measurements of sedimentation rates and sediment oxygen uptake throughout a whole year demonstrated that the benthic mineralization is governed by the sedimentation over a longer time-scale, but that seasonal imbalances do occur. A box-model of the flux of particulate organic carbon to the sediment surface is presented, and includes the relevant processes and some quantitative estimates.     

A sensitivity analysis of an ecosystem model of estuarine carbon flow

The North Inlet Marsh-Estuarine System Model (NIMES) is a 19-compartment real-time deterministic ecosystem simulation model of intrasystem carbon flow and exchange between an estuary and adjacent coastal water. A complete sensitivity analysis of this model with regard to POM, DOM and nekton annual exchange and annual system net productivity was completed and the functional relationship between these system behaviors and the perturbed parameters were determined by regression techniques. Simulated POM annual exchange between the estuary and the sea was largely controlled by offshore POM concentration, water column respiration and the gross productivity of the marsh and water column flora. Simulated DOM annual estuarine-oceanic exchange was most sensitive to perturbations in the gross productivity and biomass changes in marsh flora and water column microbial DOM uptake. Simulated nekton exchange reflected a sensitivity to migratory behavior and subtidal benthic biomass changes. System annual net productivity as simulated by the model showed a high sensitivity to all model processes which affected component primary production and respiration. From this sensitivity analysis, a scheme is developed to evaluate research needs for further model development for the North Inlet ecosystem.