Impact of heavy metals of industrial plant wastewater on benthic communities of Bizerte Lagoon (Tunisia)

ABSTRACT

The aim of this study was to assess the consequences of human impact on the characteristics of sediments heavy metal concentration, grain size and its influence on the structure of the microbial and meiofaunal community assemblages. A survey was carried out in July 2013 within six sites located in the Bizerte Lagoon (Tunisia), both downstream and upstream of industrial effluents. The highest total sediment metal concentrations were detected in stations located close to the industrial sewage discharge points. In these stations, the lowest densities of the total meiofauna (33?±?13?ind/10?cm^?2) and conversely the highest densities of cultivable bacteria that are heavy metal resistant have been reported (16?±?80.34?CFU?g^?1). Univariate (ANOVA) and multivariate (MDS/CCA) analyses demonstrate high dissimilarity (0.06) in meiofaunal and bacterial community structures between downstream and upstream industrial sewages. Furthermore, canonical correspondence analysis CCA results indicated that heavy metal sediment contamination promoted bacteria that are resistant to heavy metals, while heterotrophic bacteria supported the development of meiofauna taxa. The results highlight the importance of bacteria/meiofauna interactions, as both meiofaunal and microbial communities give indications of the ecological impact of heavy metal contamination in sediment.     

Trophic specialisation of metazoan meiofauna at the Håkon Mosby Mud Volcano: fatty acid biomarker isotope evidence

We report the results of a detailed investigation on the trophoecology of two dominant meiofaunal species at the Håkon Mosby Mud Volcano (HMMV), a deep-sea cold methane-venting seep. Analyses of fatty acids (FAs) and their stable carbon isotopes were used to determine the importance of chemosynthetic nutritional pathways for the dominant copepod species (morphologically very similar to Tisbe wilsoni) inhabiting the volcano’s centre and the abundant nematode Halomonhystera disjuncta from the surrounding microbial mats. The strong dominance of bacterial biomarkers (16:1ω7c, 18:1ω7c and 16:1ω8c) coupled with their individual light carbon isotopes signatures (δ¹³C ranging from ?52 to ?81‰) and the lack of symbiotic relationships with prokaryotes (as revealed by molecular analyses and fluorescent in situ hybridisation) indicated that chemosynthetically derived carbon constitutes the main diet of both species. However, the copepod showed a stronger reliance on the utilisation of methanotrophic bacteria and contained polyunsaturated FAs of bacterial origin (20:5ω3 and 22:6ω3 with isotope signatures δ¹³C < ?80‰). Instead, the FA profiles of H. disjuncta suggested that sulphide-oxidising bacteria constituted the main diet of this nematode. Therefore, HMMV can be regarded as a persistent deep-sea cold seep, allowing a chemosynthesis-based trophic specialisation by the dominant meiofaunal species inhabiting its sediments. The present investigation, through the determination of the fatty acid profiles, provides the first evidence for trophic specialisation of meiofauna associated with sub-habitats within a cold seep.     

The relative significance of bacteria,meio- and macrofauna on an exposed sandy beach

A survey of the intertidal biota of a sandy beach on the west coast of South Africa has shown standing stocks of macrofauna, meiofauna and bacteria of 241.23, 200.17 and 663.07 g dry wt m^-1 of shoreline respectively, an approximate biomass ratio of 1:1:3. The distribution of the macrofauna was the reverse of the usual pattern, with highest biomass occurring at the level of the current driftline. This appears to be related directly to the ready availability of food in the form of drift algae. Peak meiofaunal numbers were also found below the driftline and it is proposed that meiofaunal distribution is governed by dissolved organic matter (DOM) levels in the interstitial environment. Bacteria were abundant up to 1.2 m below the sediment surface, with the highest concentrations occurring at low tidal levels. The significance of the various biotic components in the energetics of the sandy intertidal is discussed. Turnover estimates suggest that bacteria may account for about 87% of annual production, with meiofauna and macrofauna making up 10 and 3% respectively. Despite this overwhelming importance of bacteria, the macro-and meiofauna probably play a vital role in making small organic particles available to bacteria for mineralization and in optimising conditions for microbial growth.     

Efficient size-selective bacterivory by phagotrophic nanoflagellates in aquatic ecosystems

Size-selective grazing on bacterioplankton by phagotrophic nanoflagellates was analyzed and modelled. The proposed model resembles a Monod equation (Michaelis-Menten-like expression) in which clearance rates by heterotrophic nanoflagellates depend on bacterioplnnkton biovolume. Larger bacteria were ingested faster than smaller bacteria. The proposed model was in agreement with experimental data from different authors, both from cultures and natural assemblages. Size-selective grazing efficiency was analyzed as the ability of phagotrophic nanoflagellates to discriminate between cells differing in volume by a factor of two, e.g., corresponding to dividing and nondividing bacteria. Unlike previously published models, the proposed model suggests that phagotrophic nanoflagellates could be highly effective size-selective grazers for small bacteria (<0.1 m³), which are the most common bacterial sizes in planktonic systems. However, phagotrophic nanoflagellates were unable to size-discriminate dividing and nondividing bacteria for volumes >0.1 m³. These results strongly support the hypothesis that heterotrophic nanoflagellate grazing, by discriminating between different sizes of dividing and nondividing bacteria, may actually be regulating bacterial size and growth rate in natural aquatic ecosystems.     

Patterns of micro- and meiofaunal abundance in marine sediments,measured with the adenosine triphosphate assay

Adenosine triphosphate (ATP) measurements are used to determine vertical and seasonal distributions of microorganisms and meiofauna in sediments from a 14 m-deep mud bottom in central Long Island Sound on 12 sampling dates from April 1975 to October 1976. Below the topmost 1 cm of sediment, ATP measurements can be useful in estimating and comparing standing stocks of microorganisms and meiofauna. In the top 1 cm, however, large quantities of newly settled bivalves (Yoldia limatula, Nucula annulata, and Mulinia lateralis) and juvenile polychaetes (Owenia fusiformis) in summer and fall months account for total ATP concentrations. The ATP content of individual meiofauna ranges from 1.97 ng individual copepod nauplius^-1 to 190.7±60 ng individual M. lateralis ^-1. In general, the total ATP content of individual polychaetes and bivalves is much higher than that of individuals of other groups. However, on a mg ATP per g wet or dry tissue basis, the ATP content of micro- and meiofaunal taxa are not significantly different. In addition to providing a means for comparing micro- and meiofaunal standing stocks, ATP measurements permit examination of the relative contribution of different meiofauna to the total living biomass of meiofauna in sediments. Total sediment ATP concentrations are greatest in the top 1 cm at all seasons, and decrease with increasing depth in the sediment. Annual concentrations in the topmost centimeter average 4.22 g g dry sediment^-1 and range seasonally from 1.09 to 7.64 g g dry sediment^-1. At a depth of 10 cm, values average 0.16 and range from 0.019 to 0.35 g g dry sediment^-1. High ATP concentrations in surface sediment reflect high concentrations of microorganisms and meiofauna at the sediment-water interface. The top 2 cm of sediment contain 71% of all meiofauna, with 41% occurring in the topmost cm. In general, densities are lowest in the winter and highest during the spring and summer, averaging 490 individuals 10 cm^-2, and varying from 87 to 1366 individuals 10 cm^-2. Because of wide variation in recruitment patterns of the benthos in Long Island Sound, the extremes of the range in meiofaunal densities can be observed in the same month in two different years. In order to monitor and compare standing stocks of organisms less than 1 mm in size in sediments, the ATP assay can save hours of processing time compared with alternate methods such as direct counts.     

Production of DMSP and DMS during a mesocosm study of an Emiliania huxleyi bloom: influence of bacteria and Calanus finmarchicus grazing

We investigated the influence of bacteria and metazooplankton on the production of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) during blooms of Emiliania huxleyi (Lohmann) Hay and Mohler in seawater mesocosms. The phytoplankton succession was marked by the rapid collapse of an initial Skeletonema costatum (Greville) Cleve bloom followed by a small E. huxleyi bloom. The collapse of the diatom bloom was accompanied by an increase in concentrations of dissolved DMSP (DMSP_d) and bacterial abundance and activity (as determined by the thymidine incorporation technique). The increase in bacterial activity was followed by a rapid decrease in DMSP_d concentrations which remained low for the rest of the experiment, even during the subsequent collapse of the E. huxleyi blooms. The absence of DMSP_d and DMS peaks during the declining phase of the E. huxleyi blooms was attributed to the high bacterial activity prevailing at that time. The influence of metazooplankton grazing on DMSP and DMS production was investigated by adding moderate (24 mg dry weight m^-3) and high (520 mg dry weight m^-3) concentrations of Copepodite Stage V and adults of Calanus finmarchicus to two of four filtered (200 m mesh net) enclosures during the E. huxleyi blooms. The addition of C. finmarchicus, even in high concentrations, had no apparent effect on the dynamics of E. huxleyi, suggesting that the copepods were not grazing significantly on nanophytoplankton. The addition of copepods in high concentrations favored an accumulation of chlorophyll a and particulate DMSP. These results suggest that copepods were preying on the herbivorous microzooplankton which, in turn, was controlling the biomass of nanophytoplankton. DMS production was also enhanced in the enclosure with maximum metazooplankton biomass, suggesting that the grazing of C. finmarchicus on microzooplankton containing DMSP may contribute to DMS production. These results provide strong support to the emerging idea that bacteria and metazooplankton grazing play a dominant role in determining the timing and magnitude of DMS pulses following phytoplankton blooms.     

Nitrogenous excretion by meiofauna from coral reef sediments: Mecor 5

Nitrogenous excretion rates of meiofauna from coral sediments of the Great Barrier Reef, Australia were studied at winter temperatures. Data on single taxa fitted a geometric regression: Log N excretion=-0.206+0.865 log x where log x=dry weight in g. Abundance and biomass data on meiofaunal taxa were obtained from Lizard Island, Orpheus Island and Davies Reef. Using excretion rates and biomass data, meiofauna were calculated as contributing 10.18, 12.32, and 1.60 mg N m^-2 h^-1 at the above areas respectively.Contribution No. 5 from the workshop Microbial Ecology of coral Reefs (MECOR), sponsored by the Australian Institute of Marine Sciences and the US National Science Foundation, July–August 1984     

Productivity of bacteria and microalgae and the effect of grazing by holothurians in sediments on a coral reef flat

Bacterial productivity in sandy sediments on reef flats at Lizard Island, Great Barrier Reef was determined from the rate of incorporation of tritiated thymidine into DNA. The study was conducted during January 1982 and July 1983. A small diurnal increase occurred in sediments having a dense population of microalgae. Bacterial production was 120 to 370 mg C m^-2 d^-1 in summer on reef flats, which was equivalent to 30–40% of primary production by benthic microalgae. In winter, rates of primary production by benthic microalgae and secondary production by bacteria were about one-half to one-fifth of those in summer. There was much variation in production, due to patchiness in the distribution of benthic microbes, especially microalgae. Doubling times for the bacteria in surface sediment were 1 to 2 d in summer and 4 to 16 d in winter on the reef flats. These high productivity values for bacteria indicated that a net input of organic matter to the sediment was needed to support the growth of bacteria. Sediment bacteria thus have a very important role in transforming organic matter on the reef flats. Grazing by Holothuria atra depressed both primary production and bacterial production. It was estimated that these holothurians ate about 10 to 40% of bacterial carbon produced each day in summer, and thus have an important role in the carbon cycle. Harpacticoid copepods were numerically important components of the benthic meiofaunal community and probably had a significant impact on bacterial density as grazers.     

Meiofauna densities and gastrotrich community composition in a Mediterranean sea cave

During the past 5 decades, the large-sized biota inhabiting dark marine caves has attracted the attention of many marine biologists; in contrast, studies concerning the meiofaunal organisms of these peculiar biotopes remain scanty and mostly with a taxonomic aim. In this study, the nature and abundance of meiofaunal taxa living in a Mediterranean, semi-submerged sea cave was surveyed in relation to distance from the entrance and over two different seasonal periods. Particular attention was paid to the Gastrotricha taxocene. Research was carried out in a cave along the Ionian coast of Apulia (southern Italy), the “Grotta Piccola del Ciolo” which opens for approximately 120 m on the north-eastern side of a shallow fjord and has a bottom blanketed by fine to very fine sand, occasionally rich in detritus. Quantitative samples in four replicates were collected by SCUBA diving, in November 2000 and June 2001, coring the sediment with a hand-held piston corer in three light-free sites (stations 1–3) located at increasing distances from the entrance. At each site, two additional 500-ml sediment samples were collected for an in vivo study of the Gastrotricha. Faunistic analysis found a fairly high meiobenthic diversity, identifying representatives of more than 12 major groups, with total abundances ranging from 656 ind./10 cm² (10 cm²) in November to 1,069 ind./10 cm² in June. Station 1, the closest to the entrance invariably hosted the most abundant meiofaunal community (851 ind./10 cm² in November and 1932 ind./10 cm² in June), followed by station 2 or 3 depending on the season. While nematodes and harpacticoids appear as the most abundant taxa when the cave is considered as a whole, other taxa may prevail numerically in selected stations, e.g. priapulids, which are the second most abundant taxon at station 1 (30 ind./10 cm² in November and 83 ind./10 cm² in June). Although the density of total meiofauna and that of the single groups may not be very high, the cave is interesting by virtue of the peculiarity of the hosted fauna, e.g., species and genera new to science or new to the Mediterranean Sea. Regarding the Gastrotricha, we found 16 species, accounting for 1.3–2.6% of the total meiobenthos (density = 8.4 ind./10 cm² in November and 27.4 ind./10 cm² in June). Analysis of the gastrotrich community found, particularly in June, an assemblage of taxa quite different from those found in open habitats, even at the family level; differences that are probably due to the exploitation of different food resources by animals populating the two environments, i.e. algae in the open sea versus bacteria in the caves. Results indicate that for meiofauna, as happens for macrofauna, the marine caves may represent hotspots of biodiversity and endemism; the driving forces at the base of the trophic depletion hypothesis seem to be responsible for structuring the meiofauna community inside the cave.     

Application of a 15N tracer method to the study of dissolved organic nitrogen uptake during spring and summer in Chesapeake Bay

The dissolved organic nitrogen (DON) pool in marine waters contains a diverse mixture of compounds. It is therefore difficult to accurately estimate planktonic uptake of DON using the limited number of radiolabeled compounds commercially available. We describe a method to estimate DON uptake rates using ¹⁵N-labeled DON recently released from phytoplankton. To make ¹⁵N-labeled DON, we incubated surface water with ¹⁵NH ₄ ⁺ and then isolated the DON, including any recently released DO¹⁵N, with ion retardation resin. This DON was then added to a freshly collected water sample from the same environment to quantify the rate of DON uptake. The technique was applied to investigate rates of DON uptake relative to inorganic nitrogen in the mesohaline Chesapeake Bay during May 1990 and August 1991. The May experiment took place after the spring bloom, and rates of DON uptake [ranging from 0.31 to 0.53 g-atom (g-at) Nl^-1 h^-1] often exceeded rates of NH ₄ ⁺ and NO ₃ ^- uptake combined. The rates of DON uptake at this time were higher than estimated bacterial productivity and were not correlated with bacterial abundance or bacterial productivity. They were, however, correlated with rates of NO ₃ ^- uptake. In May, we estimate that only 7 to 32% of DON uptake was a result of urea utilization. In contrast, in August, when regenerated nutrients predominate in Chesapeake Bay, rates of DON uptake (ranging from 0.14 to 0.51 g-atom Nl^-1 h^-1) were an average of 50% of the observed rates of NH ₄ ⁺ uptake. Consistent with the May experiment, rates of DON uptake were not correlated with bacterial production. A sizable fraction of DON uptake, however, appeared to be due to urea utilization; rates of urea uptake, measured independently, were equivalent to an average of 74% of the measured rates of DON uptake. These findings suggest that, during both periods of study, at least a fraction of the measured DON uptake may have been due to utilization by phytoplankton.     

Substrate-dependent bacterivory by intertidal benthic copepods

The trophic importance of bacteria to harpacticoid copepods in intertidal areas remains poorly understood, and so do the mechanisms of bacterial feeding. It is, for instance, unclear whether harpacticoids directly target bacterial cells or merely co-ingest them with substrates to which bacterial cells may be attached. Here, we investigate bacterial uptake and substrate requirement for four mud intertidal species (Microarthridion littorale, Platychelipus littoralis, Delavalia palustris and Nannopus palustris) by means of ¹³C-labeled bacteria and biomarker fatty acids (FA). Bacterial uptake strongly depended on grazing on a primary food source but bacterial ingestion rates were low, and no clear indication of copepods directly targeting bacteria was found. Delavalia was the only species that accumulated bacteria-derived FA and gained in polyunsaturated FA (PUFA) probably through bioconversion of bacteria-derived FA. In general, however, our results suggest that bacteria represent a relatively minor and low-quality food for intertidal harpacticoid copepods.     

Detritus-Bacteria-Meiofauna interactions in a seagrass bed (Posidonia oceanica) of the NW Mediterranean

The biochemical composition of the sediment organic matter, and bacterial and meiofaunal dynamics, were monitored over an annual cycle in aPosidonia oceanica bed of the NW Mediterranean to test the response of the meiofauna assemblage to fluctuations in food availability. Primary production cycles of the seagrass and its epiphytes were responsible for relatively high (compared to other Mediterranean systems) standing stocks of organic carbon in sediments (from 1.98 to 6.16 mg Cg^–1 sediment dry weight). The biopolymeric fraction of the organic matter (measured as lipids, carbohydrates, and proteins) accounted for only a small fraction (18%) of the total sedimentary organic carbon. About 25% of the biopolymeric fraction was of microphytobenthic origin. Sedimentary organic carbon was mostly refractory (56 to 84%) and probably largely not utilizable for benthic consumers. The biopolymeric fraction of the organic matter was characterized by high carbohydrate concentrations (from 0.27 to 5.31 mg g^–1 sediment dry weight in the top 2 cm) and a very low protein content (from 0.07 to 0.80 mg g^–1 sediment dry weight), which may be a limiting factor for heterotrophic metabolism in seagrass sediments. RNA and DNA concentrations of the Sediments varied significantly during the year. High RNA and DNA values occurred during the microphytobenthic bloom and in correspondence with peaks of bacterial abundance. Bacteria accounted for a small fraction of the total organic carbon (0.65%) and of the biopolymeric organic carbon (4.64%), whilst microphytobenthos accounted for 3.79% of total organic carbon and for 25.08% of the biopolymeric carbon. Bacterial abundance (from 0.8 to 5.8 × 10⁸ g^–1 sediment dry weight) responded significantly to seasonal changes of organic matter content and composition and was significantly correlated with carbohydrate concentrations. Bacteria might be, in the seagrass system, an important N storage for higher trophic levels as il accounted for 25% of the easily soluble protein. pool and contributed significantly to the total DNA pool (on average 12%). Total meiofaunal density ranged from 236 to 1858 ind. 10 cm^–2 and was significantly related, with a time lag, to changes in bacterial standing stocks indicating that microbes might represent an important resource. Bacterial abundance and biomass were also significantly related to nematode abundance. These results indicate that bacteria may play a key role in the benthic trophic     

A comparison of macrofaunal and meiofaunal distribution and standing stock across a rocky shore,with an estimate of their productivities

In 1984 and 1985 algal, macrofaunal and meiofaunal standing stocks were estimated on a exposed rocky shore along the west coast of False Bay, South Africa, using comparable, area-based sampling techniques. The shore supported a rich growth of algae, particularly in summer, when a maximum standing crop of 403 g m^-2 was recorded in the low shore. In winter, the largest component of macrofaunal biomass comprised the filter-feeding barnacle Tetraclita serrata, which attained 75 g m^-2 in the middle balanoid; but as a result of late recruitment and high mortality of this species, the summer shore was dominated by herbivorous gastropods, particularly Patella cochlear, which attained a maximum biomass of 66 g m^-2 on the low shore. Meiofaunal numbers and biomass were closely correlated to the distribution of algal turfs and associated trapped sediments. Numerically, the most important components of the meiofauna were nematodes and copepods, while the biomass was more evenly shared among foraminifera, minute gastropods, copepods and insect larvae. Numbers and biomass peaked in the lower balanoid during winter (1.9×10⁶ individuals, or 8.5 g m^-2). Macrofauna:meiofauna numbers and biomass ratios are presented for each zone and the distribution patterns discussed in relation to the conditons in each. Numerically, meiofauna exceed macrofauna by an overall ratio of 1:391, with values ranging from 1:556 in the lower balanoid to 1:18 in the Littorina zone. Macrofaunal biomass exceeds that of meiofauna in all zones by an overall ratio of 10:1, but values range from 2.1:1 in the upper balanoid to 48:1 in the middle balanoid. By incorporating turnover ratios extrapolated from the literature, mean annual productivity ratios have been calculated. These indicate that macrofauna account for 75% of total secondary production and meiofauna for 25%. Failure to incorporate meiofauna in analyses of energy flow on rocky shore ecosystems would thus lead to considerable errors. The possible trophic role of meiofauna in such systems is discussed.     

Temporal and spatial variation in bacterial production in the water column over a coral reef

Production and doubling times of the bacterial populations in the water around and over the reefs at Lizard Island, Great Barrier Reef were measured during summer and winter, 1982 and 1983. Bacterial productivity, determined from the rate of tritiated thymidine incorporation into DNA, was high over the reef flats and a Thalassia hemprichii sand flat (28 to 58 g Cl^-1 d^-1). Bacterial growth rates increased during the day and fell at night over the reef flats and seagrass bed. Growth rates were slower over the reef front and in open water. Doubling times ranged from about 2 d in the open water to about 3 h over the reef flat in summer. As numbers did not increase, grazing was probably intense on the reef flats. Growth rates were much slower in winter. The main source of organic nutrient used by the bacteria was probably mucus released following photosynthesis in the corals. The cyanobacterium Synechococcus sp. was sometimes very numerous, especially in summer when 2×10⁸ cells l^-1 were recorded in one water mass. The number of bacteria was also very high in summer, with values ranging from 1×10⁹ to 2.5×10⁹l^-1.     

Distribution patterns of meiofauna associated with a sublittoral<Emphasis Type="Italic"> Laminaria</Emphasis> bed in the Cantabrian Sea (north-eastern Atlantic)

This study deals with meiofauna associated with a sublittoral population of the kelp Laminaria ochroleuca located on the northern coast of Spain. By sampling once a year over a 4-year period, we examined patterns of faunal distribution as a function of some environmental factors at the meso-scale level (depth, and exposure to waves and surge). We also examined the relationship between L. ochroleuca abundance (as dry weight biomass and number of plants per sampling quadrat) and abundance and diversity of meiofauna. Finally, we investigated patterns of within-plant distribution (algal frond vs. algal holdfast), using also the meiofauna of the adjacent bottom as a referent to estimate the level of "phytal dependence" of the meiofauna collected on L. ochroleuca. We found that the bulk of permanent meiofauna consisted of nematodes, copepods, mites, polychaetes, tanaids and ostracods, with copepods being predominant on the fronds of the alga and nematodes in the holdfasts. The temporary meiofauna consisted of juvenile amphipods, bivalves and gastropods, together with barnacle nauplii and cyprids. Abundance and major composition of meiofaunal taxa were unrelated to both depth and hydrodynamic exposure of the sampling quadrats. However, we detected significant qualitative and quantitative faunal differences as a function of microhabitat. All meiofaunal groups were more abundant in holdfast samples than in frond and bottom samples. The gross taxonomic composition of meiofauna in bottom samples was similar to that in holdfast samples, but substantially different from that of meiofauna associated with the fronds. The L. ochroleuca holdfasts, in which dense aggregations of meiofauna can occur, appear to function as ecotone between phytal and rocky-bottom microhabitats. All together, our results suggest that the distribution of meiofauna within the Laminaria bed is mostly affected by factors operating at the microhabitat level rather than the meso-scale level.Communicated by L. Hagerman, Helsingør     

Preservation and incubation time-induced bias in tracer-aided grazing studies on meiofauna

A recent review suggests that meiofauna are important grazers of microphytobenthic primary production as well as of bacterial secondary production. The potential importance of meiofauna grazers may nevertheless have systematically been underestimated, since label leakage from chemically preserved animals has hitherto not been accounted for. Furthermore, a majority of studies have used relatively long incubation times and assumed, rather than proved, that label recycling over this period is negligible. In the present study we tested the influence of sample preservation on label retention in the marine nematode Pellioditis marina Andrassy, 1983 fed ³H-labelled bacteria. Label loss from formaldehyde-preserved specimens averaged 40% after 1 h preservation and amounted to a maximum of 85% after 24 h in formaldehyde, irrespective of formaldehyde concentration; no further leakage occurred beyond 24 h. Glutaraldehyde and ethanol yielded significantly better and poorer results, respectively, but the former fixative still yielded label losses of up to 70%. A comparison of label uptake as a function of time with observations on ingestion and defecation behaviour suggest that on time scales of hours an indication of assimilation (after correction for label leakage) rather than of ingestion is obtained. When killed with formaldehyde at room temperature, P. marina egested a significant part of its gut contents. The sources of bias identified here may have generally led to significant underestimations of true grazing rates. The cumulative effect of label leakage, prey egestion and long incubation times, each at the highest rates observed in this study, may yield as much as a 15-fold underestimation of true food consumption. Cooling samples on ice and fixation with ice-cold formaldehyde, followed by immediate freeze-preservation, and sorting of the nematodes within 2 h after thawing, gives average values for label leakage of 50%, and hence allows the application of a proximate correction factor for label losses of 2. Received: 9 July 1998 / Accepted: 23 September 1998     

Species-specific grazing rates of heterotrophic dinoflagellates in oceanic waters,measured with a dual-label radioisotope technique

A dual-isotope method was developed to measure grazing rates and food preferences of individual species of heterotrophic dinoflagellates from natural populations, collected from the Slope, Gulf Stream, and Sargasso Sea and from a transect from Iceland to New England, in 1983. The isotope method measures the grazing rates of microzooplankton which cannot be separated in natural populations on the basis of size. Tritiated-thymidine and ¹⁴C-bicarbonate were used to label natural heterotrophic and autotrophic food, respectively. Nine oceanic dinoflagellate species in the genera Protoperidinium, Podolampas, and Diplopsalis fed on both heterotrophic and autotrophic food particles with clearance rates of 0.4 to 8.0 l cell^-1 h^-1, based on ³H incorporation, and 0.0 to 28.3 l cell^-1 h^-1, based on ¹⁴C incorporation. Two dinoflagellate species, Protoperidinium ovatum and Podolampas palmipes, fed only on ³H-labelled food particles. Several species of dinoflagellates fed on bacteria (<1 m) which had been prelabelled with ³H-thymidine. The clearance rates of heterotrophic dinoflagellates and ciliates were similar and within the range of tintinnid ciliate clearance rates reported in the literature. As heterotrophic dinoflagellates and ciliates can have comparable abundances in oceanic waters, we conclude that heterotrophic dinoflagellates may have an equally important impact as microheterotrophic grazers of phytoplankton and bacteria in oceanic waters.Partially supported by a grant from the National Science Foundation, OCE-81-17744     

Uptake of dissolved organics by marine bacteria as a function of fluid motion

A mass transfer analysis predicts that fluid motion can increase the assimilation of dissolved organics by attached compared to free-living microorganisms under certain conditions. To test this we examined the effect of advective flow and fluid shear on the uptake of model compounds (leucine and glucose) by natural assemblages of heterotrophic bacteria, collected from Roosevelt Inlet, Delaware Bay (USA), in 1989. We found that [³H]leucine uptake by cells held in fluid moving at 20 to 70 m d^–1 was eight times larger than uptake by cells at a velocity of 3 m d^–1. This effect was only observed at low leucine concentrations (ca. 1 nM), when uptake was likely not saturated. When we added leucine at concentrations expected to saturate leucine uptake (ca. 11 nM), fluid motion past cells did not affect uptake. Fluid flow past bacteria did not increase [³H]glucose uptake, and laminar shear rates of 0.5 to 2.1 s^–1 did not increase either glucose or leucine uptake by suspended bacteria. These results indicate that fluid motion increases bacterial uptake of certain lowmolecular-weight dissolved organics only when the microorganism exists in an advective flow field. As predicted from a mass transfer model, fluid shear rates in natural systems are too low to affect bacterial uptake of such compounds.     

Copepod nutritional condition and pelagic production during autumn in Kosterfjorden,western Sweden

The in situ grazing rate and nutritional condition of copepods were studied during October/November 1985, by analyzing gut fluorescence (feeding), body size and lipid composition (nutritional state), and electron transport system (ETS) activity (respiration rate) of copepods from surface-and deep-water in Kosterfjorden on the Swedish west coast. These parameters were related to the physical and biological environment, as defined by light, hydrography, autotrophic and bacterial production and seston in the water column. The results show a gradual build-up of the autumn phytoplankton bloom in the uppermost meters, with a peak in total autotrophic production in mid October of ca 550 mg C m^–2 d^–1, and a bacterial net production corresponding to 15% of this. Phytoplankton exudates made up, on average, 47% of the primary production and more than 50% of this was utilized by the bacteria. Copepods occurring in the surface-water exhibited grazing rates corresponding to between 11 and 18% of their body C d^–1 and potential growth rates of 0 to 9% d^–1. Copepod populations in the surface water were composed of individuals with higher average body-weight and lower lipid-proportion than those from the deep-water.Calanus finmarchicus in the deep-water showed characters indicating diapause condition, while this was not observed forAcartia clausi. Differences in lipid content and composition indicate thatC. finmarchicus, Pseudocalanus sp. andA. clausi represent three successive points on a scale of tolerance for fluctuations in the food environment. Adult femaleMetridia longa was the only one among seven species/stages of copepods in the deep-water ( 50 m depth) that contained phytoplankton pigments.Study performed through Tjärnö Marine Biological Laboratory, University of Göteborg, S-452 00 Strömstad, Sweden     

Vertical and horizontal distribution of sub-littoral meiofauna in Algoa Bay,South Africa

Meiofauna was sampled using SCUBA along 4 transects from 5 to 30 m depth in Algoa Bay, South Africa. Substrates were well-sorted medium to fine sands with traces of sewage pollution in some areas. Meiofauna numbers recorded in the upper 10 cm of sediment were 55 to 584 10 cm^-2, but total numbers, including deeper-living animals, were 680 to 2090 10 cm^-2 at 6 stations. Longer cores showed meiofauna to be abundant down to at least 35 cm, and samples of interstitial water taken from this depth in the sediment were 7% saturated with oxygen. This is the first detailed record of meiofauna penetration into sub-littoral sand and stresses the need for long cores for quantitative work. Nematode numbers were found to be significantly related to nitrogen in the sand while interstitial harpacticoid numbers were related to median particle diameters, which determine pore space. Effects of sewage were slight and only one station showed notable enrichment, increased meiofaunal numbers and extreme dominance by nematodes.