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.     

Energy budgest,growth and filtration rates in Mytilus edulis at different algal concentrations

Growth of Mytilus edulis L. was measured in aquaria with through-flowing sea water at different levels of constant algal concentrations. The amount of food and oxygen consumed by the mussels were measured over given periods as well as the changes in dry organic weight during the same periods. From these parameters it was possible to make simple energy budgets and to compare the estimated growth with actual growth, and, further, to determine growth efficiences at different food levels. Energy budgets were made for mussels grown at algal concentrations of 0, 1.6×10³, 3.0×10³ and 26.0×10³ Phaeodactylum tricornutum cells x ml^-1. The estimated growth was found to be close to actual growth at algal concentrations above maintenance level and the net growth efficiency was found to be between 18% (3.0×10³ cells x ml^-1) and 61% (26×10³ cells x ml^-1). It has been shown that the filtration rate is independent of algal concentrations between about 1.5×10³ to 30×10³ P. tricornutum cells x ml^-1. Outside this range a decrease in filtration rate was noticed.     

Particle retention and selection by larvae and spat of Ostrea edulis in algal suspensions

Grazing rates and electivity indices of larvae and spat of Ostrea edulis L. were, measured and examined in relation to certain physical parameters using a flow-through system. Retention and size-selection were determined for the major particle sizes present in cultures of Isochrysis galbana Parke, an alga used frequently as food for bivalves. Cultures of the algae Dunaliella tertiolecta Butcher and Phaeodactylum tricornutum Bohlin were used as sources of particle suspensions of various sizes and shapes, respectively. While increases in flow rate caused increased grazing, the mode of selection of I. galbana particles remained constant. Filtration rate, F _f was related to body size, W, by the general allometric equation R _f =aW ^b,while particle-size preference in suspensions of I. galbana by both larvae and spat of O. edulis was independent of W. Grazing rates increased with temperature to an optimum temperature, which was related to the acclimation temperature. Increases above this optimum caused a reduction in feeding activity. No significant change in particle size-preference in the I. galbana suspension with temperature was observed. Grazing rates and selection were dependent, however, on particle number and volume. Both larvae and spat displayed maximum retention at optimum particle concentrations which tended to decrease with increasing particle size. Variations in cell shape of P. tricornutum had no measurable effect on selectivity by O. edulis.     

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.     

Adjustment of the components of energy balance in the gastropod Crepidula fornicata in response to thermal acclimation

Synchronous measurements were made of the routine rate of oxygen consumption and the clearance rate of Phaeodactylum tricornutum at different exposure temperatures by specimens of the suspension-feeding gastropod Crepidula fornicata which had been acclimated to temperatures between 10° and 25°C. The results show that the cost of activity (l O₂ consumed h^-1/ml seawater cleared h^-1) increases dramatically in individuals exposed to short-term increases of temperature up to 30°C, especially in limpets acclimated to 10°C. The process of thermal acclimation, however, results in two compensatory adjustments in energy expenditure and uptake which profoundly affect the energetics of water transport. Firstly, the routine oxygen consumption shows lateral translation of the rate-temperature curve which results in the maintenance of a relatively uniform energy expenditure despite an increase in acclimation temperature from 10° to 25°C. Secondly, because of the form of the rate-temperature curve for filtration by C. fornicata, lateral translation in response to warm acclimation results in an increase in the maximal clearance rate. Lateral translation of the rate-temperature curves for feeding rates and for oxygen consumption in response to thermal acclimation may thus be linked to maintain a balance between energy gain and expenditure. In this way, the greatly increased cost of activity which would occur with increase of temperature in the absence of acclimation is evaded. The minimal maintenance energy requirement, and hence the greatest scope for growth and reproduction, is then adjusted to coincide with temperatures prevailing in the environment.     

Growth and lipid composition of scallop juveniles, Placopecten magellanicus, fed the flagellate Isochrysis galbana with varying lipid composition and the diatom Chaetoceros muelleri

The effect of feeding the flagellate Isochrysis galbana (Parke; clone T-Iso) of modified lipid composition on the growth and lipid composition of juvenile scallops [Placopecten magellanicus (Gmelin)] was investigated in the spring of 1993. I. galbana grown in 85-liter cage culture turbidostats under conditions of nitrogen limitation had a significantly higher total lipid content than when grown under nutrient-replete conditions. This was due mainly to a doubling in the amount of less unsaturated triacylglycerol in the cells. The concentrations of methyl and ethyl ketones were also greater in nitrogen-limited cells. Diets of nitrogen-limited I. galbana and nutrient-replete I. galbana grown in continuous and semi-continuous cultures were compared. Scallop juveniles were batch fed daily, and measurements of ingestion were determined. Samples of juveniles were removed periodically for determination of organic weight. The juveniles did not grow when fed nitrogen-limited or nutrient-replete I. galbana alone; however, when each diet was supplemented with 20% of the diatom Chaetoceros muelleri (Lemm.), there was a significant increase in growth in the juveniles receiving the nitrogen-limited I. galbana compared with juveniles on other diets. In comparison with I. galbana,  C. muelleri provided a rich source of carbohydrates and the essential fatty acid 20:4ω6. This study shows the importance of providing optimal dietary levels of ω3 and ω6 polyunsaturated fatty acids, as well as less unsaturated fatty acids and carbohydrates. Received: 29 September 1997 / Accepted: 2 October 1998     

Growth and competition of the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana. II. Light limitation

The marine diatoms Phaeodactylum tricornutum (Bohlin) and Thalassiosira pseudonana (Hasle and Heimdal) were grown under both continous illumination and a 14 h light: 10 h dark cycle at light intensities ranging from 1.53×10^-4 to 2.95×10^-1 ly min^-1. Under both photoperiods, T. pseudonana exhibited higher division rates than P. tricornutum at high light intensities, but the reverse was true at all light intensities <3×10^-3 ly min^-1. Comparison of these results with available data on light-limited growth of other planktonic algae suggests that P. tricornutum may be unusually efficient at maintaining its cell division rate at low light intensity. This efficiency may contribute substantially to its success in turbid, nutrient-enriched mass algal culture systems, the only environments in which it is known to attain great numbers.Contribution No. 4086 from the Woods Hole Oceanographic Institution.     

Grazing of Acartia hudsonica (A. clausi) on Skeletonema costatum in Narragansett Bay (USA): Influence of food concentration and temperature

Grazing experiments were performed with temperatureacclimated Acartia hudsonica fed the diatom Skeletonema costatum in concentrations ranging from 50 to 3×10⁴ cell ml^-1 at 5°, 10° and 15°C. The ingestion data were best fit by an Ivlev equation. Feeding threshold values of 39 and 59 cells ml^-1 were not significantly different from zero; however, filtration rates were depressed at low food concentrations. Maximum filtration rates increased exponentially with temperature, reaching a maximum with copepods collected at 14°–15°C, and then declining. Both the increase in ingestion rate with increasing food concentration and the maximum ingestion rate were significantly greater as experimental temperature was increased. Maximum ingestion rates were reached at concentrations greater than 6×10³ cells ml^-1. Percent of body carbon ingested per day at 5 g C L^-1 increased from 1.5% at 5°C to 6.7% at 15°C. At 500 g C L^-1, the ingestion increased from 84% (5°C) to 660% (15°C). Percent of body nitrogen at 0.5 g N L^-1 increased from 0.6% per day at 5°C to 2.5% per day at 15°C. At 50 g N L^-1, the ingestion was 42% body nitrogen at 5°C and 250% at 15°C. The influence of grazing by A. hudsonica on phytoplankton in Narragansett Bay, USA was estimated for 1972–1977. The percent of standing stock removed by grazing rarely exceeded 5% per day except during the late spring when S. costatum growth becomes nutrient limited and higher temperatures favor the rapid population growth of A. hudsonica.     

Filter-feeding ethology of benthic invertebrates (ascidians). V. Influence of temperature on pumping,filtration and digestion rates and rhythms in Phallusia mamillata

The influence of temperature has been studied simultaneously on the pumping, filtration, and digestion rates of Phallusia mammillata (Cuvier, 1815). Eighteen experiments were made between 7° and 25°C on 5 individuals. The average velocities of the water current varied between 3.37 and 9.65 cm sec^-1 (maximum 34.90 cm sec^-1). No recognizable rhythm emerged; pumping was continuous except at 7°C, where it soon ceased. Above 20°C, the curves were irregular and reflected the high sensitivity of the ascidian. The pumping rate was highest at 15°C (mean=5,788 ml h^-1 g^-1 dry weight of organs). At 10°C, the mean was 3,560; at 20°C, 2,629 ml h^-1 g^-1 dry weight of organs. At 20°C, the coefficients of variation displayed higher values, indicating a more irregular pumping at this temperature. Although there was no filtration rhythm, the variability of the results was higher at 20°C and above. As for pumping, maximum values were observed at 15°C (mean=4,286 ml h^-1 g^-1 dry weight of organs) decreasing with lower and higher temperatures, such decreases being more marked at the higher temperatures. Means were 352 ml h^-1 g^-1 dry weight of organs at 7°C; 2,935 at 10°C; 1,995 at 20°C; 973 at 25°C. The mean temperature coefficients for the filtration rates were: Q₁₀ for 7° to 15°C=11.86, Q₁₀ for 10° to 20°C=0,66, Q₁₀ for 15° to 25°C=0.22. The filtering efficiency was fairly constant throughout an experiment; the pumping and filtration curves were in fact almost parallel. The filtering efficiency of the branchial sac was high (75 to 85%), with constant values at 10° and 15°C; it became smaller (59%) at 20°C, with a higher coefficient of variation. The digestion rate also displayed maximum values at 15°C (mean=5.47 mg of albumin equivalent 24 h^-1 g^-1 dry weight of organs). It was lower at 10°C (mean=3.60 mg) and reached its minimum at 20°C (mean=1.71 mg). The higher temperature affected the percentage of food utilization, which showed smaller values at 20°C (59%) than at 10°C (89%) and 15°C (87%).     

Food limitation stimulates metamorphosis of competent larvae and alters postmetamorphic growth rate in the marine prosobranch gastropodCrepidula fornicata

The effects of food limitation on growth rates and survival of marine invertebrate larvae have been studied for many years. Far less is known about how food limitation during the larval stage influences length of larval life or postmetamorphic performance. This paper documents the effects of food limitation during larval development (1) on how long the larvae ofCrepidula fornicata (L.) can delay metamorphosis in the laboratory after they have become competent to metamorphose and (2) on postmetamorphic growth rate. To assess the magnitude of nutritional stress imposed by different food concentrations, we measured growth rates (as changes in shell length and ash-free dry weight) for larvae reared in either 0.45-m filtered seawater or at phytoplankton concentrations (Isoehrysis galbana, clone T-ISO) of 1 × l0³, 1 × 10⁴, or 1.8 × 10⁵ cells ml^–1. Larvae increased both shell length and biomass at 1 × 10⁴ cells ml^–1, although significantly more slowly than at the highest food concentration. Larvae did not significantly increase (p > 0.10) mean shell length in filtered seawater or at a phytoplankton concentration of only 1 × 10³ cells ml^–1, and in fact lost weight under these conditions. To assess the influence of food limitation on the ability of competent individuals to postpone metamorphosis, larvae were first reared to metamorphic competence on a high food concentration ofI. galbana (1.8 × 10⁵ cells ml^–1). When at least 80% of subsampled larvae were competent to metamorphose, as assessed by the numbers of indlviduals metamorphosing in response to elevated K⁺ concentration in seawater, remaining larvae were transferred either to 0.45-m filtered seawater or to suspensions of reduced phytoplankton concentration (1 × 10³, 1 × 10⁴, or 5 × 10⁴ cells ml^–1), or were maintained at 1.8 × 10⁵ cells ml^–1. All larvae were monitored daily for metamorphosis. Individuals that metamorphosed in each food treatment were transferred to high ration conditions (1.8 × 10⁵ tells ml^–1) for four additional days to monitor postmetamorphic growth. Competent larvae responded to all food-limiting conditions by metamorphosing precociously, typically 1 wk or more before larvae metamorphosed when maintained at the highest food ration. Surprisingly, juveniles reared at full ration grew more slowly if they had spent 2 or 3 d under food-limiting conditions as competent larvae. The data show that a rapid decline in phytoplankton concentration during the larval development ofC. fornicata stimulates metamorphosis, foreshortening the larval dispersal period, and may also reduce the ability of postmetamorphic individuals to grow rapidly even when food concentrations increase.     

Development and metamorphosis of the planktotrophic larvae of Rostanga pulchra (Mollusca: Nudibranchia)

Rostanga pulchra MacFarland, a small (1 to 2 cm) dorid nudibranch, lays an average of 7000 eggs in the laboratory during a period of 30 days in the summer. The veligers hatch 15 to 16 days after oviposition and it takes another 35 to 40 days to become competent for metamorphosis at a temperature of 10° to 15°C. Larval cultures were maintained initially at a concentration of 500 veligers per 100 ml of filtered sea water (antibiotics added). During the planktotrophic phase of development, the veliger grows from 150 to 300 m in shell length. Although the veligers are generalists in their food preference, the best result (faster growth) was achieved by feeding them with a combination of Monochrysis lutheri and Isochrysis galbana. The concentration of food cells was kept at 10⁴ cells per ml of culture media and was supplied every 2 to 3 days. A veliger which is competent to metamorphose is identifiable morphologically by its propodium, eyespots, rhinophores, and spiculated dorsal papillae. The entire metamorphic process lasts 24 h when a suitable substrate such as the food sponge Ophlitaspongia pennata is provided. The competent veliger is able to delay metamorphosis for at least 3 weeks. Juveniles were kept in the laboratory for 70 days and, during this period, grew to a length of 4.5 mm.     

Shell growth response of mussels (Mytilus edulis) exposed to toxic microalgae

Mussels (Mytilus edulis) were exposed to the algaeAlexandrium ostenfeldii, Chrysochromulina polylepis, Gyrodinium aureolum, Gymnodinium galatheanum andHeterosigma akashiwo for 24 h; significant reductions in growth rate, as compared to the control, were observed after exposure toA. ostenfeldii, C. polylepis, G. aureolum andG. galatheanum at initial concentrations of 4.5 × 10⁶, 110 × 10⁶, 9 × 10⁶ and 120 × 10⁶ cells l^–1, respectively. Exposure to high initial concentrations of the non-toxic algaeTetraselmis suecica (174 × 10⁶ cells l^–1) andIsochrysis galbana (610 × 10⁶ cells l^–1) showed no adverse effect on growth rate. When mussels with reduced growth were transferred to clean seawater, they recovered to > 90% of control growth within 2 to 4 d. Exposure to algal filtrates of the toxic algal cultures produced no reduction in growth rate.     

Assessing the toxicity of chemical compounds associated with marine land-based fish farms: The use of mini-scale microalgal toxicity tests

Many chemicals that are currently used in aquaculture have not been evaluated with regard to their specific effects on the aquatic environment. In the present study, the toxic effects of several chemicals associated with land-based marine fish farming activities were assessed using two species of marine microalgae (Phaeodactylum tricornutum and Isochrysis galbana). Mini-scale toxicity tests were performed with six antibiotics (amoxicillin, ampicillin, flumequine, oxytetracycline, streptomycin and sulfadiazine) and two disinfectants (formaldehyde and hypochlorite). Amoxicillin and streptomycin did not exert toxic effects. Sulfadiazine was the most toxic chemical; the EC₅₀ values were 0.11 mg/L and 1.44 mg/L for P. tricornutum and I. galbana respectively. As expected, the disinfectants displayed high toxicity, and P. tricornutum was particularly sensitive to these compounds. Although the differences in microalgal sensitivity depended on the chemical considered, both species were highly sensitive to most of the compounds tested. We recommend the inclusion of mini-scale microalgal toxicity tests in environmental risk assessment (ERA) and environmental monitoring plans because they are cost-effective and rapid.     

Chemical composition of dietary species of marine unicellular algae and rotifers with emphasis on fatty acids

The lipid profiles of a few species of marine unicellular algae and yeast were studied with emphasis on fatty acids as part of a search for the nutritional value of plankton to the diet of marine fish larvae commonly used in marine hatcheries. The general proximate chemistry of rotifers was closely related to the proximate chemistry of the diet organism, exhibiting a higher content of protein and carbohydrate and a lower content of lipid. Major lipids in all algae, yeast and rotifers comprised mono-, di- and tri-glycerides and polar lipids. The algae Chaetoceros gracilis Schutt, Isochrysis galbana Parke and their respective algaefed rotifers exhibited higher amounts of neutral lipids, consisting mainly of cyclic and branched polyunsaturated components. Fatty acid composition of the algae was species-specific, with the highest ratio of polyethylenic to saturated and monoethylenic acid in I. galbana and Phaeodactylum tricornutum Bohlin, and the highest content (15%) of n-3 highly unsaturated fatty acids in Nannochloropsis salina and P. tricornutum. A closely mirrored distribution of the fatty acids, but with a lower amount of n-3 highly unsaturated fatty acids, was present in the respective algae-fed rotifers. Comparison of the fatty acid spectrum of Artemia sp. and Euterpina acutifrons grown in the laboratory on I. galbana with zooplankton samples of E. acutifrons and Oitona nana collected from the sea showed a higher concentration of docosahexaenoic acid (22:6 n-3) in the naturally collected sample. The results indicate that the efficacy of the food algae C. gracilis and I. galbana in increasing the survival of fish larvae in marine hatcheries is not obvious on the sole basis of fatty acid composition.     

Carbon conversion and metabolic rate in two marine sponges

The carbon metabolism of two marine sponges, Haliclona oculata and Dysidea avara, has been studied using a ¹³C isotope pulse-chase approach. The sponges were fed ¹³C-labeled diatoms (Skeletonema costatum) for 8 h and they took up between 75 and 85%. At different times, sponges were sampled for total ¹³C enrichment, and fatty acid (FA) composition and ¹³C enrichment. Algal biomarkers present in the sponges were highly labeled after feeding but their labeling levels decreased until none was left 10 days after enrichment. The sponge-specific FAs incorporated ¹³C label already during the first day and the amount of ¹³C label inside these FAs kept increasing until 3 weeks after labeling. The algal-derived carbon captured by the sponges during the 8-h feeding period was thus partly respired and partly metabolized during the weeks following. Apparently, sponges are able to capture enough food during short periods to sustain longer-term metabolism. The change of carbon metabolic rate of fatty acid synthesis due to mechanical damage of sponge tissue was studied by feeding sponges with ¹³C isotope–labeled diatom (Pheaodactylum tricornutum) either after or before damaging and tracing back the ¹³C content in the damaged and healthy tissue. The filtration and respiration in both sponges responded quickly to damage. The rate of respiration in H. oculata reduced immediately after damage, but returned to its initial level after 6 h. The ¹³C data revealed that H. oculata has a higher metabolic rate in the tips where growth occurs compared to the rest of the tissue and that the metabolic rate is increased after damage of the tissue. For D. avara, no differences were found between damaged and non-damaged tissue. However, the filtration rate decreased directly after damage.     

Relationship between filtration activity and food availability in the Mediterranean mussel <Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis>

The filtration activity of the Mediterranean mussel, Mytilus galloprovincialis, was assessed under different concentrations and compositions of seston by using a new automated image acquisition and analysis system. This approach allowed for frequent and simultaneous measurements of valve gape and exhalant siphon area. Filtration rates were measured through clearance measurements whereas pumping rates were measured using hot-film probes. The average filtration rate (17.5 l g h⁻¹ DW⁻¹ for a 0.36 g DW mussel) recorded during the present study was higher than those available for Mytilus edulis when standardized to flesh dry weight but almost equivalent (17.5 l h⁻¹ g DW⁻¹ for a 53 mm shell length mussel) to those rates when standardized to shell length. Immediately after the addition of algal cells (Isochrysis galbana; 4.5 μm in size), valve gape, exhalant siphon area and filtration rate increased quickly as mussels reached their maximum filtration activity. These three parameters then gradually decreased until complete closure of the shell. The algal cell concentration inducing this transition was close to 800 cells ml⁻¹ and 0.5 μg Chl a l⁻¹. When algal concentration was maintained above this threshold by successive algal additions, both valve gape and exhalant siphon area remained maximal. Temporal changes in the exhalant siphon area were continuous as opposed to those of valve gape. Therefore, despite the significant correlation between these two parameters, valves and siphon were sometimes dissociated due to a reduction of the area or even a closure of the exhalant siphon while the valves remained open. The velocity of exhaled water tended to be constant irrespective of exhalant siphon area and thus pumping rates were a linear function of exhalant siphon area. Consequently, reductions in exhalant siphon area and pumping rate were almost similar in M. galloprovincialis. Our results thus clearly support the hypothesis that exhalant siphon area constitutes a better proxy of pumping rate than valve gape as already suggested for Mytilus edulis. Finally, the high filtration rates measured during the present study together with the high concentrations of inorganic matter (> 40 mg DW l⁻¹) requested to alter those rates suggest that the studied mussels were well adapted to oligotrophic waters featuring strong hydrodynamism and frequent sediment resuspension events.     

Growth and competition of the marine diatoms Phaeodactylum tricornutum and Thalassiosira pseudonana. I. Nutrient effects

Two marine diatoms, Phaeodactylum tricornutum (Bohlin) and Thalassiosira pseudonana (Hasle and Heimdal), were grown both separately and together in batch cultures on a mixture of waste water and seawater enriched with different components of f medium. At 17°C, the maximum division rates of the two species were statistically indistinguishable. The waste water-seawater mixture used proved to have insufficient Si, relative to N and P, for the growth of T. pseudonana, which requires approximately 5x10^-14 g-at Si cell^-1 to divide at a maximum rate. P. tricornutum, on the other hand, although capable of taking up nearly 9x10^-15 g-at Si cell^-1, could sustain maximum rates of division with 4.3x10^-18 g-at Si cell^-1 or less. No allelopathic interaction between the two species could be detected. We conclude that P. tricornutum enjoys a considerable competitive advantage over T. pseudonana in a waste water-seawater-based mariculture system that is not supplemented with Si. Although Si proved necessary for T. pseudonana to complete more successfully with the other diatom, the presence of excess amounts of Si is not necessarily sufficient for the maintenance of T. pseudonana in mixed continuous culture with P. tricornutum: other factors, such as light-related or photoperiod-related growth response, are believed to determine the ultimate outcome of competition between these algae in light-limited continuous culture.Contribution No. 3999, from the Woods Hole Oceanographic Institution.Communicated by M.R. Tripp, Newark     

Comparison of larval development and growth of the sea cucumberActinopyga echinites: Ovary-induced ova and DTT-induced ova

AdultActinopyga echinites (Jaeger) were collected from northern Taiwan in September 1989. Oocytes were induced to mature by bathing them in ovary juice (ovary-induced ova) or in 10^–2 M dithiothreitol (DTT-induced ova). The percentage of germinal vesicle breakdown (GVBD) increased from 0.4 to 6.4% in the former treatment and to 84% in the latter. After artificial fertilization, the embryos were cultured in seawater (35 S) at 25 to 28°C. Larvae were fed with the algaIsochysis aff.galbana at 10^4–5 cells/ml. Larvae from ovary-induced ova developed faster (18 d to the doliolaria stage) and grew to a larger size (1.13 mm length) than those from DTT-induced ova (20 d to the doliolaria stage and 0.62 mm in length). On the twelfth day, larvae from ovary-induced ova bear lipid spheres. The number of spheres is positively correlated with larval size. Lipid spheres may provide nutrient reserves for larvae during metamorphosis.     

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.     

Anaerobic heat production of bivalves (Polymesoda caroliniana andModiolus demissus) in relation to temperature,body size,and duration of anoxia

Anaerobic heat-production rates of two co-occurring species of estuarine bivalves (a clam and a mussel) were measured with double-twin heat-flow calorimeters, one at 20°C, the other at 30°C. There is no significant difference between the two species in metabolic rates. There is evidence of initial aerobic metabolism in some individuals, as shown by high initial rates exponentially decreasing with time, while others had fluctuating but stable average metabolic activity from the beginning. During aerobic as well as anaerobic metabolism, the bivalves showed rhythmic periods of activity and quiescence. The two species differed in their rhythmic pattern of active and resting metabolism. In the case ofPolymesoda caroliniana, periods of resting metabolism tend to be longer and periods of active metabolism shorter at 30°C than at 20°C. There is a similarity between thermograms ofModiolus demissus at 20° and 30°C. Following acute temperature changes from 5° to 20° and 30°C, the bivalves showed stable metabolic rates in a matter of hours. The stabilized average rates [pooled averages for both species of 1.34×10^-4 (standard error of the mean=0.17×10^-4) W g^-1 dry weight of tissue at 20°C and 2.10×10^-4 (SE=0.20×10^-4) W g^-1 at 30°C] signify a temperature coefficient (Q₁₀) of 1.56 between 20° and 30°C, or partial temperature acclimation. Subtracting heat production as a result of physical activity, i.e., considering only resting metabolism, the corresponding means and standard errors of the means are 1.24×10^-4 and 0.14×10^-4 W g^-1 at 20°C and 1.91×10^-4 and 0.077×10^-4 W g^-1 at 30°C. Anaerobic heat production rate at 20°C is proportional to body size (r=0.84, 9 degrees of freedom, DF). ForM. demissus, measured anaerobic heat production is on the order of 7.5% of the level of aerobic respiration reported in the literature.