Diet of intertidal bivalves in the Ría de Arosa (NW Spain): evidence from stable C and N isotope analysis

We used natural abundance measurements of stable carbon and nitrogen isotopes and a short-term ¹⁵N-enrichment experiment to explore the importance of river-derived and autochthonous sources to the diet of intertidal suspension-feeding bivalves in the Ría de Arosa, an oceanic-dominated system in northwest Spain. Highly seasonal and intense phytoplankton "bloom" events occur in this estuary. However, relatively low phytoplankton standing crop and productivity are present during much of the year, suggesting that food sources other than phytoplankton must support bivalves during these periods. Bivalves (Cerastoderma edule, Tapes decussatus, Mytilus galloprovincialis) were sampled quarterly at four intertidal stations along the length of the estuary with increasing distance from the Ulla River. Muscle tissue was consistently enriched in ¹³C and ¹⁵N relative to stomach contents. The stable C isotope values of bivalves indicated that river-derived inputs were, at most, a minor component of diet. These values suggested the potential importance of benthic microalgae to bivalve diet much of the year and the incorporation of ¹⁵N label applied to the sediment surface into stomach contents and muscle tissue in a field experiment supported this conclusion. Other labeled components of the microbenthos such as bacteria could also have been ingested. ¹³C-depleted values of bivalves in May coincided with elevated offshore chlorophyll a concentrations, suggesting the increased use of a phytoplankton source during the spring. We propose that intertidal suspension-feeding bivalves in this estuary use primarily resuspended microbenthos during periods of low phytoplankton concentration, but that phytoplankton increases in relative dietary importance during bloom events.Communicated by P.W. Sammarco, Chauvin     

Filtration rate,using a new indirect technique,in thirteen species of suspension-feeding bivalves

A method for determining filtration rates in undisturbed suspension-feeding bivalves is described. Concentrations of particulate matter in the water collected in the inhalant (C _i) and exhalant (C _e) currents were estimated with an electronic particle counter. The clearance was calculated as , where Fl=flow rate through the tube collecting exhalant water. Only above critical levels of water flow (Fl) were clearances representative of filtration rates. At 10° to 13°C, the filtration rates (F, 1 h^-1) within one or two orders of magnitude of dry weight (w, g), in Cardium echinatum L., C. edule L., Mytilus edulis L., Modiolus modiolus (L.) and Arctica islandica (L.) followed the allometric equations: 4.22w ^0.62, 11.60w ^0.70, 7.45w ^0.66, 6.00w ^0.75 and 5.55w ^0.62, respectively. Five species of bivalves [Spisula subtruncata (da Costa), Hiatella striata (Fleuriau), Cultellus pellucidus (Pennant), Mya arenaria L. and Venerupis pullastra (Montagu)] filtered with the same rates as individuals of Cardium echinatum and A. islandica of equivalent soft weight. In Pecten furtivus and P. opercularis filtration rates were about twice the rates measured in individuals of Mytilus edulis of comparable body weight. The gill area in M. edulis increases with size at the same rate as the filtration rate.     

An improved automatic recording apparatus for determining the filtration rate of Mytilus edulis as a function of size and algal concentration

An automatic recording apparatus for measuring the filtration rate in suspension-feeding bivalves is described. The concentration of algae in the experimental medium is kept constant throughout each experiment by addition of Phaeodactylum tricornutum from a chemostat. Within the range of body size 5.7 to 283 mg (W=dry weight of tissues), the filtration rate (F=ml min^-1) at 15°C in Mytilus edulis L. follows the allometric equation F=0.85 W ^0.72. Within the concentrations 0.18 to 0.70 mg algal dry weight l^-1, the filtration rate in mussels of 132 mg dry flesh weight ranges from 33.1 to 41.0 ml min^-1. At 0.18 mg algal dry weight l^-1 the mussels filter continuously for 20 h, with a high constant rate that presumably represents the water transport capacity under optimal laboratory conditions.     

Calorimetric studies on the energy metabolism of an infaunal bivalve,Abra tenuis,under normoxia,hypoxia and anoxia

Direct calorimetry was employed to measure the energy metabolism of infaunal bivalves, Abra tenuis, collected from a tidal lagoon in the Fleet, southern England, in June 1989, at various oxygen partial pressures. A significant anaerobic component (i.e., 20% of total metabolic rate) was detected under normoxia, presumably brought about by the intermittent ventilatory activity of this bivalve under these conditions. Under hypoxia (2.3 to 10 kPa, or 11 to 48% of full air saturation), however, the energy metabolism was maintained fully aerobic; the measured heat equivalent of oxygen uptake was not significantly different from the theoretical ranges for fully aerobic catabolism. Under anoxia, the rate of heat dissipation was reduced to 5–6% of the normoxic rate of heat dissipation. This conserves energy expenditure and would thus increase resistance of A. tenuis to anoxia or emersion. Physiological compensation by A. tenuis under conditions of declining oxygen tension involved a marked increase in ventilation rate. Comparison between fed and starved individuals indicated that costly physiological processes, such as digestion, absorption and growth declined at 10 and 5 kPa and were arrested at P_{O 2} (oxygen partial pressure) levels below 2.3 kPa. The present study provides evidence that there are no major differences between the metabolic responses of epifaunal suspension-feeding (eg. Mytilus edulis) and infaunal deposit-feeding (eg. A. tenuis) bivalves when exposed to environmental hypoxic stress.     

The influence of suspension density and temperature on the filtration rate of Hiatella arctica

The rate of filtering Phaeodactylum tricornutum and Isochrysis galbana was measured in Hiatella arctica (L.) by the indirect suspension depletion method monitored by optical density measurement. The filtration rate of H. arctica was found to be 1.412×10^–2 l/h/g wet weight at a temperature of 15°C when fed with P. tricornutum, at average cell concentrations up to 3.5×10⁶ cells/ml. The filtration rate dropped almost to zero when the concentration of P. tricornutum reached 11×10⁶ cells/ml. The filtration rate of I. galbana diminished at a much lower cell concentration of 1×10⁶ cells/ml, and almost ceased at 3 to 4×10⁶ cells/ml. In mixed cultures of I. galbana and P. tricornutum, the filtration rate ratio was 0.37 to 1.00, and this was believed to be due to a proportion of the smaller former cells passing through the ostia. However, when resuspended in sea water, I. galbana cells were taken at a rate slightly less than P. tricornutum. The medium in which the I. galbana cells had been grown was inhibitory to the filtering activity of H. arctica, since, when cells of either alga were resuspended in the medium, the filtration rate was considerably reduced. No inhibitory factor existed in either of the original nutrient media. Hence, the importance of using low cell concentrations and of eliminating any inhibitory metabolic products when measuring filtration rates of bivalves is stressed. H. arctica shows a typical activity temperature eurve for a boreo-arctic species, with a steady rise from 0°C to a maximum between 15° and 17°C, and a sharp fall in activity to about zero at 25°C. The rates of filtration of various species at temperatures approaching the optimum were compared after allowance was made for fall in filtration rate with increasing body weight. The results suggested that the Mytilacea had the highest filtration rates and that H. arctica possesses one of the lowest filtration rates recorded.     

Localization of infaunal prey by the sea star<Emphasis Type="Italic"> Leptasterias polaris</Emphasis>

We performed field and laboratory studies to investigate how large adult Leptasterias polaris detect and locate their major prey, large infaunal bivalves, in the sediment bottom community. A field survey using SCUBA diving showed that 95% of the locations where L. polaris dug into the sediment bottom were over bivalves and this success rate was much greater than if digging was done at random (22%). Furthermore, when sea stars were provided with a low density of randomly distributed prey in a laboratory arena, they dug exclusively in locations where a clam had been buried. These observations indicated that L. polaris locates infaunal prey prior to investing energy into digging. Studies in a laboratory flow tank showed that L. polaris readily detected and moved towards its preferred prey Ensis directus whereas its responses to less preferred prey Mya truncata and Spisula polynyma were much weaker. The degree to which it oriented towards these three common prey seemed to reflect potential energy intake relative to foraging costs (which likely increase with the depth of the different prey) and risks from interactions with other carnivores (which are greatest when feeding on large prey). This is the first study to clearly demonstrate that sea stars use prey odours to locate infaunal prey.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-004-1497-1Communicated by R.J. Thompson, St. Johns     

Über den Einfluß der Nahrungskonzentration und anderer Faktoren auf Filtrierleistung und Nahrungsausnutzung der Muscheln Arctica islandica und Modiolus modiolus

Filtration rates and the extent of phagocytosed food particles were determined in the offshore lamellibranchs Artica islandica and Modiolus modiolus in relation to particle concentration, body size and temperature. Pure cultures of the algae Chlamydomonas sp. and Dunaliella sp. were used as food. A new method for determining filtration rates was developed by modifying the classical indirect method. The concentration of the experimental medium (100%) was kept constant to ±1%. Whenever the bivalves removed algae from the medium, additional algae were added and the filtration rate of the bivalves expressed in terms of percentage amount of algae added per unit time. The concentration of the experimental medium was measured continuously by a flow colorimeter. By keeping the concentration constant, filtration rates could be determined even in relation to different definite concentrations and over long periods of time. The amount of phagocytosed food was measured by employing the biuret-method (algae cells ingested minus algae cells in faeces). Filtration rates vary continuously. As a rule, however, during a period of 24 h, two phases of high food consumption alternate with two phases of low food consumption during which the mussels' activities are almost exclusively occupied by food digestion. Filtration rate and amount of phagocytosed algae increase with increasing body size. Specimens of A. islandica with a body length of 33 to 83 mm filter between 0.7 to 71/h (30–280 mg dry weight of algae/24 h) and phagocytose 21 to 122 mg dry weight of algae during a period of 24 h. The extent of food utilization declines from 75 to 43% with increasing body size. In M. modiolus of 40 to 88 mm body length, the corresponding values of filtration rate and amount of phagocytosed algae range between 0.5 and 2.5 l/h (20–100 mg dry weight of algae) and 17 to 90 mg dry weight of algae, respectively; the percentage of food utilization does not vary much and lies near 87%. Filtration rate and amount of phagocytosed algae follow the allometric equation y=a·x ^b. In this equation, y represents the filtration rate (or the amount of phagocytosed algae), a the specific capacity of a mussel of 1 g soft parts (wet weight), x the wet weight of the bivalves' soft parts, and b the specific form of relationship between body size and filtration rate (or the amount of phagocytosed algae). The values obtained for b lie within a range which indicates that the filtration rate (or the amount of phagocytosed algae) is sometimes more or less proportional to body surface area, sometimes to body weight. Temperature coefficients for the filtration rate are in Arctica islandica Q₁₀ (4°–14°C)=2.05 and Q₁₀ (10°–20°C)=1.23, in Modiolus modiolus Q₁₀ (4°–14°C)=2.33 and Q₁₀ (10°–20°C)=1.63. In A. islandica, temperature coefficients for the amount of phagocytosed algae amount to Q₁₀ (4°–14°C)=2.15 and Q₁₀ (10°–20°C)=1.55, in M. modiolus to Q₁₀ (4°–14°C)=2.54 and Q₁₀ (10°–20°C)=1.92. Upon a temperature decrease from 12° to 4°C, filtration rate and amount of phagocytosed algae are reduced to 50%. At the increasing concentrations of 10×10⁶, 20×10⁶ and 40×10⁶ cells of Chlamydomonas/l offered, filtration rates of both mollusc species decrease at the ratios 3:2:1. At 12°C, pseudofaeces production occurs in both species in a suspension of 40×10⁶, at 20°C in 60×10⁶ cells of Chlamydomonas/l. At 12°C and 10–20×10⁶ cells of Chlamydomonas/l, the maximum amount of algae is phagocytosed. At 40×10⁶ cells/l, the amount of phagocytosed cells is reduced by 26% as a consequence of low filtration rates and intensive production of pseudofaeces. At 20°C and 20–50×10⁶ cells of Chlamydomonas/l, the maximum amount of algae is sieved out and phagocytosed; the concentration of 10×10⁶ cells/l is too low and cannot be compensated for by increased activity of the molluscs. With increasing temperatures, the amount of suspended matter, allowing higher rates of filtration and food utilization, shifts toward higher particle concentrations; but at each temperature a threshold exists, above which increase in particle density is not followed by increase in the amount of particles ingested. Based on theoretical considerations and facts known from literature, 7 different levels of food concentration are distinguishable. Experiments with Chlamydomonas sp. and Dunaliella sp. used as food, reveal the combined influence of particle concentration and particle size on filtration rate. Supplementary experiments with Mytilus edulis resulted in filtration rates similar to those obtained for M. modiolus, whereas, experiments with Cardium edule, Mya arenaria, Mya truncata and Venerupis pullastra revealed low filtration rates. These species, inhabiting waters with high seston contents, seem to be adapted to higher food concentrations, and unable to compensate for low concentrations by higher filtration activities. Adaptation to higher food concentrations makes it possible to ingest large amounts of particles even at low filtration rates. Suspension feeding bivalves are subdivided into four groups on the basis of their different food filtration behaviour.     

Behavior and growth ofMercenaria mercenaria during simulated storm events

Experiments were conducted in April–August 1989 on juvenileMercenaria mercenaria (L.) in an oscillatory water tunnel to simulate resuspension of bottom sediments by waves and to determine the effects of shortterm storm events on particle ingestion, pseudofeces production, and shell growth. Juveniles (mean length = 19.2 mm) were subjected to identical concentrations of algae in both low-flow, gentle waves (maximum velocity = 7 cm s^–1) and high-velocity storm waves (maximum velocity = 22 cm s^–1). Suspended sediment levels reached 193 mg 1^–1 at 1 cm above the bed during storms. Shell growth decreased by a maximum of 38% during the storm when levels of phytoplankton were high (average cell concentration = 43 × 10⁶ cells 1^–1), and by 18% when phytoplankton levels were low (av cell conc = 6 × 10⁶ cells 1^–1). Orientation of clam siphons was not related to flow direction. Significantly more pseudofeces were produced when the clams were subjected to increased sediment resuspension under waves, and in troughs of sand ripples. The size of sediment grains ingested did not vary significantly among the flow treatments. The decrease in shell growth during storms may be due to a reduction in filtration rate coupled with a decrease in net energy gained from filtration due to costs of pseudofeces production. The magnitude of the decrease seems to be related to concentration of algae, water temperature, age of clams and sediment transport mode (bed load or suspended load). Thus, the interpretation of growth increments must be made in the context of these environmental variables.     

Influence of body size on the uptake,depuration, and bioaccumulation of polychlorinated biphenyl congeners by Baltic Sea blue mussels,Mytilus edulis

The present study was designed to examine the influence of body size on the bioaccumulation of polychlorinated biphenyl (PCB) congeners by Baltic Sea blue mussels, Mytilus edulis L. This was done, firstly, by establishing the relationship (as a power function: PCB tissue conc = a tissue dry wt^b) between tissue concentration and body weight for seven PCB congeners (International Union of Pure and Applied Chemistry No. 52, 101, 105, 118, 138, 153, and 180) in field sampled mussels; and, secondly, by assessing the influence of body weight on the uptake clearance coefficients, the depuration rate coefficients and the calculated bioaccumulation factors (BAFs) of three ¹⁴C-labelled PCB congeners (IUPAC No. 31, 49, 153) in mechanistic kinetic experiments. Both the background tissue concentrations of PCB 138, PCB 153 and PCB 180 and the predicted BAF values in the kinetic experiments correlated negatively with body weight (b=-0.17 and-0.31, respectively). Of the two kinetic rate coefficients examined, only the uptake clearance rate showed weight dependency (b=-0.32, i.e. negative correlation with body weight), whereas depuration rates were unaffectd by body weight. Uptake clearance rates and BAFs adjusted for body weight increased with the octanol/water partition coefficient (K_ow) of the congener, whereas depuration rates dectreased with K_ow. These observations suggest that size-dependent bioaccumulation of hydrophobic organic contaminants (HOCs) in suspension-feeding bivalves is driven by size-related differences in uptake rate, although several other mechanisms may also affect bioaccumulation in natural mussel beds (e.g. sizerelated differences in lipid content, production, and contaminant exposure). From an ecotoxicological perspective size-dependent bioaccumulation implies not only that variability due to body size differences has to be dealt with in experimental designs, but also that several ecological factors such as size-specific predation and shifts in population structure may affect HOC cycling by dense populations of bivalve suspensionfeeders.     

Particle-Size-dependent maximum grazing rates for Temora longicornis fed natural particle assemblages

The hypothesis that lower retention efficiencies of filter-feeding copepods for small particles should result in different ingestion rate versus food concentration curves for different-sized foods was tested using Temora longicornis (Müller) fed natural phytoplankton. The copepods were fed different natural phytoplankton assemblages, which varied in their species and size distribution. Volume ingestion rates were an asymptotic function of food concentration, with maximum ingestion rates measured at food concentrations exceeding 5 to 10x 10⁶ m³ ml^-1, which were less than those occurring in the natural waters in which the copepods and phytoplankton were collected. Maximum volume ingestion rates increased linearly by a factor of 3.5, as the diameter of the particle forming the peak in the food size distribution increased fron 5 m (primarily microflagellates) to 30 m (mostly large diatoms). These results suggest that natural and pollutant-induced size reductions in natural phytoplankton could markedly decrease the volume of food consumed by filter-feeding copepods.Contribution No. 243 of the Marine Sciences Research Center     

Clearance rates of the great scallop (<Emphasis Type="Italic">Pecten maximus</Emphasis>) and blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) at low natural seston concentrations

Great scallop, Pecten maximus, and blue mussel, Mytilus edulis, clearance rate (CR) responses to low natural seston concentrations were investigated in the laboratory to study (1) short-term CR variations in individual bivalves exposed to a single low seston diet, and (2) seasonal variations in average CR responses of bivalve cohorts to natural environmental variations. On a short temporal scale, mean CR response of both species to 0.06 μg L⁻¹ chlorophyll a (Chl a) and 0.23 mg L⁻¹ suspended particulate matter (SPM) remained constant despite large intra-individual fluctuations in CR. In the seasonal study, cohorts of each species were exposed to four seston treatments consisting of ambient and diluted natural seston that ranged in mean concentration from 0.15 to 0.43 mg L⁻¹ SPM, 0.01 to 0.88 μg L⁻¹ Chl a, 36 to 131 μg L⁻¹ particulate organic carbon and 0.019 to 0.330 mm³ L⁻¹ particle volume. Although food abundance in all treatments was low, the nutritional quality of the seston was relatively high (e.g., mean particulate organic content ranged from 68 to 75%). Under these low seston conditions, a high percentage of P. maximus (81–98%) and M. edulis (67–97%) actively cleared particles at mean rates between 9 and 12 and between 4 and 6 L g⁻¹ h⁻¹, respectively. For both species, minimum mean CR values were obtained for animals exposed to the lowest seston concentrations. Within treatments, P. maximus showed a greater degree of seasonality in CR than M. edulis, which fed at a relatively constant rate despite seasonal changes in food and temperature. P. maximus showed a non-linear CR response to increasing Chl a levels, with rates increasing to a maximum at approximately 0.4 μg L⁻¹ Chl a and then decreasing as food quantity continued to increase. Mean CR of M. edulis also peaked at a similar concentration, but remained high and stable as the food supply continued to increase and as temperatures varied between 4.6 and 19.6°C. The results show that P. maximus and M. edulis from a low seston environment, do not stop suspension-feeding at very low seston quantities; a result that contradicts previous conclusions on the suspension-feeding behavior of bivalve mollusks and which is pertinent to interpreting the biogeographic distribution of bivalve mollusks and site suitability for aquaculture.     

Interaction and impacts of two introduced species on a soft-sediment marine assemblage in SE Tasmania

Introduced species are having major impacts in terrestrial, freshwater and marine ecosystems world-wide. It is increasingly recognised that effects of multiple species often cannot be predicted from the effect of each species alone, due to complex interactions, but most investigations of invasion impacts have examined only one non-native species at a time and have not addressed the interactive effects of multiple species. We conducted a field experiment to compare the individual and combined effects of two introduced marine predators, the northern Pacific seastar Asterias amurensis and the European green crab Carcinus maenas, on a soft-sediment invertebrate assemblage in Tasmania. Spatial overlap in the distribution of these invaders is just beginning in Tasmania, and appears imminent as their respective ranges expand, suggesting a strong overlap in food resources will result from the shared proclivity for bivalve prey. A. amurensis and C. maenas provide good models to test the interaction between multiple introduced predators, because they leave clear predator-specific traces of their predatory activity for a number of common prey taxa (bivalves and gastropods). Our experiments demonstrate that both predators had a major effect on the abundance of bivalves, reducing populations of the commercial bivalves Fulvia tenuicostata and Katelysia rhytiphora. The interaction between C. maenas and A. amurensis appears to be one of resource competition, resulting in partitioning of bivalves according to size between predators, with A. amurensis consuming the large and C. maenas the small bivalves. At a large spatial scale, we predict that the combined effect on bivalves may be greater than that due to each predator alone simply because their combined distribution is likely to cover a broader range of habitats. At a smaller scale, in the shallow subtidal, where spatial overlap is expected to be most extensive, our results indicate the individual effects of each predator are likely to be modified in the presence of the other as densities increase. These results further highlight the need to consider the interactive effects of introduced species, especially with continued increases in the number of established invasions.Communicated by M.S. Johnson, Crawley     

Bacterial nitrification activity directly associated with isolated benthic marine animals

Potential nitrification rates (PNR) directly associated with isolated marine macrobenthic invertebrates were measured for a range of benthic epifaunal and infaunal species (bivalves, gastropods, polychaetes and crustaceans) collected from the Sacca di Goro, Po River delta, Italy. In the case of the filter-feeding bivalves, Tapes philippinarum and Mytilus galloprovicialis the PNR associated with the shell surfaces and dissected animal tissues (gills, siphons and residual tissue) were determined separately, in order to assess the distribution of the nitrifier populations. Significant PNR was found associated with all the tested macrofaunal species with activities ranging between 12 and 2,250 nmol ind.^–1 day^–1 and specific activities between 150 and 18,400 nmol g^–1 dry weight day^–1. However, no simple relationships were observed between PNR and the animals taxonomic or functional group, or with animal comportment (infaunal or epifaunal) or size class, indicating that more complex interactions may regulate the degree of colonisation of the animals by nitrifier populations. Incubations of shells alone and dissected tissues of the bivalves T. philippinarum and M. galloprovicialis demonstrated that approximately 50% of the total PNR activity was associated with the shell surfaces and 50% with the internal animal tissues, with the highest specific activities of 950 and 1,970 nmol g^–1 dry weight day^–1 determined for the gills of T. philippinarum and M. galloprovicialis, respectively. Thus, specific relationships may exist between the nitrifiers and their animal hosts. Overall, our data indicate that the macrofaunal stimulation of nitrification and/or coupled nitrification–denitrification observed in previous studies may not be solely due to the animals burrow walls serving as sites for nitrification, but also to the fact that the internal and external surfaces of the animals themselves are also colonised by nitrifying bacteria. Tentative calculations based on reported animal densities in the Sacca di Goro and the determined PNRs indicate that animal-associated nitrifier populations could contribute significantly to overall nitrification rates in situ, although further experiments are required to determine to what extent the potential rates measured in this study are realised under in situ conditions.Communicated by R. Cattaneo-Vietti, Genova     

The filtration rate of Mytilus edulis and its dependence on algal concentration,measured by a continuous automatic recording apparatus

A new apparatus for long-term, continuous automatic measurements of filtration rates in suspension-feeding organisms is described. As the concentration of algae in the experimental medium is diminished by the filter-feeding activity of the experimental animals, algal suspension is automatically added, thus keeping the algal concentration constant. In this way, accurate determinations of filtration rates in relation to particle concentration are made possible. For determination of filtration rates in the common mussel Mytilus edulis L., individuals of different body size (shell length 8.5 to 56.5 mm) were used. Within the range of 10x10⁶ to 40x10⁶ cells of Dunaliella marina/l, mussels of the same body size filter-out approximately the same amount of algae at high or low concentrations. A low algal concentration is counterbalanced by a corresponding higher filtration rate. Within the range of body size (W=dry weight of tissues) and algal concentrations used, the filtration rate (F) follows the general allometric equation F=a·W ^b, where a and b are constants at specific experimental conditions. At a temperature of 12 °C, the values obtained for a are 2410 at a concentration of 20x10⁶, and 1313 at a concentration of 40x10⁶ Dunaliella cells/l; correspondingly, the filtration rates of a mussel of 1 g dry-tissue weight are 2410 ml/h and 1313 ml/h. b, the slope of the regression line (0.73 to 0.74), is independent of algal concentration. However, examination of all known measurements reveals that, most probably, the general allometric equation is an oversimplification; in large individuals there is a more pronounced decrease in filtration rate. The relationship between filtration rate, body size of mussels, and algal concentrations used is discussed.This work was made possible through a research grant from the Deutsche Forschungsgemeinschaft in connection with the program Litoralforschung — Abwässer in Küstennähe.     

Accumulation of metals by marine picoplankton

Pumping rates in undisturbed suspension-feeding bivalves were measured with a thermistor microflowmeter. Pumping rates are related to the dry weight of Clinocardium nuttallii, Macoma nasuta, Mytilus californianus and Chlamys hastata as the following equations: P=0.85 W^0.84, P=0.23 W^0.855, P=2.216 W^0.715, and P=2.419 W^0.943 respectively. Pumping rate and gill area increase at the same rate in C. nuttallii, M. californianus and C. hastata. The pumping rate per unit gill area is highest for C. nuttallii (approximately 0.14 cm³/s/cm²), with intermediate values for M. californianus (0.08 cm³/s/cm²) and C. hastata (0.11 cm³/s/cm²); the lowest rate was found in the deposit-feeder M. nasuta (0.0057 to 0.0089 cm³/s/cm²), which also has the smallest gill to body weight. C. nuttallii has the relatively smallest gill of the suspension feeders and the highest pumping rate per unit gill area. The absolute pumping rates, however, were highest for the filibranchs M. californianus and C. hastata. Indications are given that these differences are the results of different fluid-mechanical/ecological strategies to minimize the cost-benefit ratio.Contribution No. 132 of the Marine Science and Maritime Studies Center, Northeastern University     

Effects of suspended sediments on egg production of the calanoid copepod Acartia tonsa

The estuarine copepod Acartia tonsa was collected on several occasions between 4 April and 14 August 1985 from Terrebonne Bay, Louisiana (29°08N; 90°36W) and the effects in its diet of suspended sediments, collected from the same area, were measured at five different concentrations of sediment (100 to 1 000 ppm) and six phytoplankton concentrations (500 to 13 000 cells ml^-1 Thalassiosira weissflogii). Egg production rate was used as an index of diet quality. At low phytoplankton concentrations (500 cells ml^-1), and at intermediate phytoplankton concentrations (2 000 cells ml^-1) for previously starved copepods, egg production was reduced by up to 40% at a sediment concentration of 250 ppm and further reduced at higher sediment concentrations. At higher food concentrations (4 000 to 13 000 cells ml^-1), suspended sediment had no effect on egg production rates at sediment concentrations up to 500 ppm. Rates were reduced only at the highest sediment concentration of 1 000 ppm. Under most natural conditions, suspended sediment would not significantly affect egg production rates in A. tonsa.     

Description and use of an improved method for determining estuarine zooplankton grazing rates on phytoplankton

A method of rapidly determining zooplankton grazing rates on natural mixed phytoplankton populations using ¹⁴C is described. The method simplifies the design of grazing experiments as the grazing time can be kept short enough to prevent recycling of the isotope, and growth of the phytoplankton substrate. Very high specific activity, ¹⁴C-labelled phytoplankton concentrated either by centrifugation or sieving, may be used either as the sole grazing substrate, or as a tracer in natural mixed phytoplankton. Zooplankton, confined in glass jars at either ambient, or higher than ambient concentrations, are permitted to feed on the phytoplankton for periods of 30 min and 2 h, and are then separated by sieving. The zooplankton community grazing rate, or, if the samples are sorted into species, the individual species grazing rates, can be determined after scintillation counting of the zooplankton. The rate of appearance of ¹⁴C-labelled phytoplankton in the zooplankton is an estimate of the grazing rate, and the slope of the line joining the grazing rates at various phytoplankton concentrations gives an estimate of the grazing rate constant for the zooplankton population. The method provides a quick way of obtaining both zooplankton population, and individual species grazing rates on natural mixed phytoplankton. In two experiments, labelled phytoplankton was used as the sole grazing substrate in concentrations ranging between 0.4 and 5 times ambient levels. Grazing rate constants, for net-caught zooplankton concentrated to 46 times (Experiment 1) and 28 times (Experiment, 2) ambient estuarine levels were-0.14and-0.12 of the phytoplankton standing stock per day, respectively. There was a linear increase in the amount of phytoplankton grazed with an increase in phytoplankton concentration up to four times ambient phytoplankton levels. When tracer amounts of labelled phytoplankton were added to samples containing both phytoplankton and zooplankton at ambient concentrations the grazing rate constants were-0.28 and-0.42 of the phytoplankton standing stock per day. We conclude that zooplankton grazing was the major control factor of phytoplankton population size during October–November 1975 in South West Arm, Port Hacking, near Sydney, Australia.     

Particle sorting and formation and elimination of pseudofaeces in the bivalves <Emphasis Type="Italic">Mulinia edulis</Emphasis> (siphonate) and <Emphasis Type="Italic">Mytilus chilensis</Emphasis> (asiphonate)

Mulinia edulis and Mytilus chilensis are suspension-feeding bivalves with homorhabdic gills that live in different sedimentary habitats in the lower and upper intertidal, respectively, in Yaldad Bay, Chile. They are faced with different suspended particle size distributions when feeding, and both eliminate most of the inorganic particles by pseudofaeces production. This study used histology, scanning and transmission electron microscopy, and video endoscopy to compare particle processing on the labial palps and the mechanisms of particle sorting, acceptance, and rejection in the two species. In both species, disaggregation of mucus-bound particles occurs on the plicate surface of the palps. Particles destined for ingestion pass anteriorly from crest to crest and reach the mouth via the palp acceptance tract. Those destined for rejection enter the troughs between the plicae and move ventrally to the palp rejection tract. The palps manipulate the pseudofaeces into a mucous ball, which is transferred to the mantle rejection tract. In Mulinia edulis (an infaunal, siphonate mactrid), the pseudofaeces are stored in a chamber at the base of the inhalant siphon until expelled by intermittent contraction of the siphon wall. In contrast, Mytilus chilensis (an epifaunal, non-siphonate mytilid) releases pseudofaeces continuously when submerged.     

In situ water motion and nutrient uptake by the giant kelp Macrocystis pyrifera

Rates of NO ₃ ^- uptake by individual blades of Macrocystis pyrifera (L.) C. Agardh were measured at different flow rates in the laboratory. Dissolution rates of hemispherical, plaster buttons attached to the blade surface provided a relative measure of flow rates over blades used in uptake experiments and also over intact blades of adult kelp plants in situ (Laguna Beach, California, USA; 1981). Laboratory results indicated that uptake was saturated at a flow rate equivalent to 2.5 cm s^-1 current velocity. Flow rates over intact blades in situ always exceeded this uptake saturation level. Wave surge and movement of plant surfaces relative to the surrounding water provided sufficient flow to saturate uptake, even in a dense kelp canopy during low-current and calm sea-state conditions.     

Relationship between phytoplankton cell size and the rate of orthophosphate uptake: in situ observations of an estuarine population

Orthophosphate uptake by a natural estuarine phytoplankton population was estimated using two methods: (1) ³²P uptake experiments in which filters of different pore sizes were used to separate plankton size-fractions; (2) ³³P autoradiography of phytoplankton cells. Results of the first method showed that plankton cells larger than 5 m were responsible for 2% of the total orthophosphate uptake rate. 98% of the total uptake rate occurred in plankton composed mostly of bacteria, which passed the 5 m screen and were retained by the 0.45 m pore-size filter. There was no orthophosphate absorption by particulates in a biologically inhibited control containing iodoacetic acid. Orthophosphate uptake rates of individual phytoplankton species were obtained using ³³P autoradiography. The sum of these individual rates was very close to the estimated rate of uptake by particulates larger than 5 m in the ³²P labelling experiment. Generally, smaller cells were found to have a faster uptake rate per m³ biomass than larger cells. Although the nannoplankton constituted only about 21% of the total algal biomass, the rate of phosphate uptake by the nannoplankton was 75% of the total phytoplankton uptake rate. Results of the plankton autoradiography showed that the phosphate uptake rate per unit biomass is a power function of the surface: volume ratio of a cell; the relationship is expressed by the equation Y=2x10^-11 X ^1.7, where Y is gP m^-3 h^-1 and X is the surface: volume ratio. These results lend support to the hypothesis that smaller cells have a competitive advantage by having faster nutrient uptake rates.