Some relationships between growth,metabolism and food in the mussel Mytilus edulis

Measurements of ingested ration, assimilated ration and metabolic rate in Mytilus edulis L. of different sizes have been integrated to provide an estimate of energy balance, which in turn describes the physiological state of the animal. These data allow the empirical determination of growth efficiency and ration. Growth efficiency increases hyperbolically with increasing ingested ration to reach a maximum, after which efficiency decreases as ration is further increased. The optimum ration for efficient growth increases with increasing weight of the mussel; maximum growth efficiency decreases with increasing body weight.     

Lead uptake from sea water and food,and lead loss in the common mussel Mytilus edulis

Common mussels, Mytilus edulis (shell length 19 to 21 mm, average dry weight 30 mg) were maintained for 6 weeks in sea water containing different concentrations of lead (0.005 to 5 mg · l^-1). The lead concentration in the mussels' whole soft parts was analysed at different times during the experiment. A constant rate of lead uptake, linearly dependent on the lead concentration of the medium, was observed. Thus, the temporal change of the concentration factor is also linear (regression coefficient 149.9 daily). Rate of lead loss, measured after transferring the mussels into natural sea water, is linearly dependent on the original lead concentration in the soft parts. Rates of uptake and loss in large mussels (shell length 45 to 55 mm, average dry weight 750 mg) are less than those in small mussels (shell length 19 to 21 mm, average dry weight 30 mg). During a much more extended experimental period, adjustment to a steady state is expected to occur; rates of lead uptake and loss are then non-linear. Lead uptake by individual organs (kidney, gills, adductor muscle, digestive gland, foot, mantle with gonads) of large M. edulis (shell length 45 to 55 mm, average dry weight 750 mg) was analysed in 2 test series. In the test series medium, the mussels were kept in a seawater medium containing 0.01 mg. Pb.l^-1. In the test series food, the mussels were kept in natural sea water but fed with the green algae Dunaliella marina containing lead (approximately 600 g.g^-1 dry weight). The lead quantity given per mussel per day was about 2 g in both test series. Within 35 days, the mussels of test series medium took up 29% of the total amount of lead given, those of test series food took up 23.5%. In all organs, lead concentration increased, but rates of uptake differed; the kidney displayed by far the highest rate of uptake. With these physiological properties M. edulis is an ideal indicator organism for lead pollution in the marine environment. A biologic calibration curve, the relationship between lead concentration in the mussels' whole soft parts at equilibrium and lead concentration in sea water, is presented.This paper forms part of a doctoral thesis in biology at Hamburg University     

Digestive rhythms in the mussel Mytilus edulis

Mytilus edulis L., collected from a mid-tide level on the shore, showed rhythmic changes in mantle fluid pH, crystalline style pH, style length and total protein, and in the amylase activity in the digestive gland. These changes were correlated with the changes in tidal height. Style size may be related to extracellular digestion in the stomach. Style size and amylase content of the style were not significantly correlated with each other. The changes in amylase activity in the digestive gland confirmed the existence of a tidal rhythm for intracellular digestion in M. edulis.     

Relationships between carbon and cadmium uptake in Mytilus edulis

The influence of six different food rations on carbon assimilation and net incorporation, and on shell and soft body growth in Mytilus edulis was determined over a 75-d period, starting in 1981. Using a triple tracer technique, interrelationships between carbon net incorporation (C-14) and cadmium accumulation from food (Cd-109) and seawater (Cd-115m) were investigated. With decreasing food rations there was an exponential increase of the carbon assimilation efficiency from 35 to 76% while the net incorporation efficiency increased approximately linearly, ranging from 15 to 32% at the highest and lowest food supply levels respectively. The carbon assimilation efficiency adapted to new conditions within 1 to 2 d while the adaptation of the net incorporation efficiency took about one month. Shells kept growing, even when the soft body lost weight. Within a 42-d elimination period, when the soft body lost 58% of the incorporated carbon, nothing was released from the shells. Cadmium accumulation from food was closely interrelated with carbon net incorporation and exhibited the same uptake kinetics. No homogeneity was found between carbon net incorporation and cadmium accumulation from seawater, and between the elimination of carbon and cadmium.     

Relationships between effects of environmental temperature and seston on growth and mortality of Mytilus edulis in a temperate northern estuary

The growth and mortality of experimentally-rafted Mytilus edulis L. of known age at 7 locations in a northern estuary (Damariscotta River, Maine, USA) were related to environmental temperatures and to the presence or absence of various potential food sources. All particles were regarded as potential food substrates. Growth decreased appreciably at sites where water temperatures exceeded 20°C, but mussels survived a wide range of elevated temperature exposures, ranging from 0 to 149 degree-days in excess of 20°C. The maximum temperature was 25°C. Mortalities of mussels at all sites but did affect the extent of mortality, which increased abruptly in late summer, when water temperatures were declining. Differences in degree-days of exposure to elevated temperatures did not influence the timing of mortality, which occurred synchronously at all times, but did affect the extent of mortality, which ranged from 35 to 90%. The period of high mortality was preceded by a rapid decline in phytoplankton standing crop. Total particle concentrations decreased during this period, but the shift toward larger particles suggests that there was little, if any, decrease in total volume of material in the seston, at least through August. It is suggested that living phytoplankton provided the critical energy source for these mussels, and that the mortalities were caused by rapidly reduced ration at a time of metabolic stress. The role of temperature and the possible role of the gametogenic cycle are discussed. In addition, the feeding efficiency of M. edulis may have decreased during this period due to an increase in mean particle diameter. Chlorophyll was divided into a nannoplankton fraction (<-20 m) and a larger fraction (>20 m). A smaller size criterion for this distinction is proposed for future studies. The importance of nannoplankton to the primary production of this estuary and the role of nannoplankton and nannoplankton-sized particles in the diet of mussels in nature are discussed.     

Copper-induced differential mortality in the mussel Mytilus edulis

Copper toxicity experiments were performed with 3 000 mussels (Mytilus edulis L.) from a population that is a mixture of individuals originating in the Kattegat and in the Baltic. Subsequently, the electrophoretic patterns of the enzymes phosphoglucose isomerase (PGI) and phosphogluco mutase (PGM) were analysed in the dead and surviving mussels in order to detect possible differential mortality. Significant differences in allele frequencies between dead and surviving mussels were found for PGI. The results are discussed in relation to genotypic differences in the PGI and in relation to the composite structure of the investigated population. It is concluded that the differential mortality is due to genotypic differences in the PGI locus and/or to different susceptibility to copper by mussels originating in the two geographic areas.     

Relationship between cadmium concentrations in seawater and those in the mussel Mytilus edulis

Various forms of regression analysis are presented which show that there is a significant equilibrium relationship between total recoverable cadmium in seawater and its concentration in the mussel Mytilus edulis (P<0.001). The calculations show that the concentration of cadmium in seawater should not exceed 0.20 g l^-1 if the mussel is not to reach a cadmium concentration of 2 mg kg^-1 wet weight: a value frequently used as a food standard for human consumption. Further, when a cadmium value of 2 mg kg^-1 wet weight is reached, the concentration factor by mussels for cadmium from seawater is 9 950.     

Formation of metallothioneins in relation to accumulation of cadmium in the common mussel Mytilus edulis

Accumulation rates of Cd have been studied in the common mussel Mytilus edulis L. under different laboratory conditions. Semi-static and through-flow experiments were carried out at Cd concentrations of 200 ppb and 50/100 ppb in the water, respectively. A linear uptake of Cd throughout the experimental periods of 36 or 124 d was found in all experiments. The accumulation rate was 3.10 ppm d^-1 at 50 ppb in the through-flow water experiment, 4.1 ppm d^-1 in starved and 6.6 ppm d^-1 in fed mussels in the semi-static experiments with 200 ppb Cd in the water. The Cd-accumulation rate in various fractions of soft parts decreased in the order: body>mantle>muscles. Elution patterns obtained from gel-filtrations showed an increasing amount of Cd-binding proteins (metallothioneins) when the body burden of Cd increased. The Cd content in the body fraction of mussels transferred to clean water for 42 d after first being exposed to Cd during 124 d decreased from 564 ppm on a dry weight basis to 417 ppm while the fraction of Cd bound to metallothioneins rose from 22 to 78% during the same period.     

Hydrocarbon uptake and loss by the mussel Mytilus edulis

The dynamics of accumulation and elimination of hydrocarbons by the blue mussel Mytilus edulis were studied in a continuous-flow system. Mussels were exposed for as long as 41 days to 200 – 400 g/l of diesel fuel adsorbed on kaolin particles. Hydrocarbons were accumulated in the tissues in excess of 1000 times the exposure levels. Upon termination of dosing, the mussels exhibited a rather rapid loss of hydrocarbons for the first 15 to 20 days (biological half-life=2.7 to 3.5 days). Subsequently, however, elimination was reduced to a minimum and a considerable fraction of the hydrocarbons could be recovered from the tissues after as long as 32 days of depuration. The mussels exhibited definite signs of physiological stress due to chronic exposure to diesel fuel, although recovery was rapid upon termination of dosing. It is concluded that mussels could be utilized as a test organism for monitoring long-term hydrocarbon pollution in marine waters. The implications for the mussel culture industry are discussed.     

Accumulation of cadmium and bioenergetics in the mussel Mytilus edulis

Accumulation rates of cadmium, the amount of food ingested and assimilated, the amount of oxygen consumed and changes in dry flesh weight have been measured in Mytilus edulis L. exposed to 0, 10 and 100 ppb cadmium for 17 d in aquaria with seawater flowing continously and at constant algal concentration. The accumulation rates were linear at 10 and 100 ppb, amounting to 0.58 and 8.89 ppm d^-1, respectively. Body loads up to 150 ppm caused no effects on either clearance, ingestion, assimilation, respiration, or growth. High net growth efficiencies between 55–59% were obtained, indicating near optimal experimental conditions. It is suggested that the setup and experimental procedure provide an excellent tool in the study of accumulation and sublethal effects of environmental pollutants in suspension feeding bivalves.     

Influence of marine microalgal metabolites on the feeding behavior of the blue mussel Mytilus edulis

Effects of microalgal ectocrines on the feeding behavior of the blue mussel Mytilus edulis L., collected from Delaware Bay, Lewes, USA, were studied. Unialgal cultures of six species of marine microalgae were grown to the end of their exponential growth phase. Cells were removed by gentle filtration, leaving dissolved microalgal ectocrines in culture filtrates. Filtrates were then tested in filtration rate and particle selection bioassays. In filtration rate bioassays, dissolved ectocrines were delivered to mussels in a flow-through apparatus and removal of beads (6 m) used to determine rates. Filtration rates were significantly reduced by dissolved ectocrines from two of the microalgal species tested (Olisthodiscus luteus Carter and Dunaliella tertiolecta Butcher). None of the microalgal filtrates stimulated filtration rates of mussels. In particle selection bioassays, ectocrines were adsorbed onto either reverse phase (lipophilic) or normal phase (hydrophilic) microparticles (10 m) and delivered to mussels with an equal concentration of control treated particles. Mussels showed significant selection, either rejection or preferential ingestion, for ectocrine treated spheres depending upon microalgal species and sphere type used. This research shows that pre-ingestive chemical cues from microalgae can influence mussel feeding behavior. We suggest that epicellular ectocrines are more important in mediating this behavior than dissolved ectocrines.     

Relationship between body zinc concentration and allometric growth measurements in the mussel Mytilus edulis

The frequency distribution of shell lengths in the mussels Mytilus edulis (L.) taken from a site in the Tyne Estuary (UK) in summer, 1980, typically displayed a bimodal shape, with a small peak in the 8 to 20 mm size range and a much broader peak in the 22 to 50 mm region. A collection of mussels was made consisting of two groups, one from each of the above size peaks. The group of larger mussels had a significantly higher mean zinc concentration than the group of smaller mussels, mainly because a few large individuals had very high zinc concentrations. The ratios of shell width:height, width:length and length:height were also significantly higher in larger mussels. However, the ratio of flesh dry wt:shell dry wt (flesh condition) was lower in larger individuals. The whole soft tissue zinc concentration was positively correlated with width:height and width:length, but negatively correlated with flesh wt:shell wt. It was concluded that allometric ratios may provide an attractive alternative to simple size characteristics as a basis for trace metal determinations, particularly in a comparison of metal levels in mussel populations from widely differing habitats where absolute size is a poor indicator of age or growth rate.Contribution No. 1221 of the Center for Estuarine and Environmental Studies of the University of Maryland     

Match/mismatch between the Mytilus edulis larval supply and seston quality: effect on recruitment

We considered Cushing's match/mismatch theory in a heterotrophic environment and hypothesized that settlement and recruitment success in blue mussel are higher when the food supply is rich in polyunsaturated and essential fatty acids (PUFA/EFA). To test this hypothesis, we monitored larval development as well as fatty acid composition in trophic resources during two successive reproductive seasons. The decoupling we found between the presence of competent larvae in the water column and settlement rates strongly suggests that metamorphosis is delayed until conditions are suitable. In both years, the major mussel settlement peak was synchronized with a phytoplanktonic pulse rich in EFA, consisting of a large autotrophic bloom in 2007 and a short but substantial peak of picoeukaryotes in 2008. These results suggest a "trophic settlement trigger" that indirectly affects recruitment by strongly improving the settlement rate. Despite similar larval settlement rates during both years, the lower 2007 recruitment likely resulted from a mismatch with a high lipid-quality trophic resource. The seasonal trophic conditions differed greatly between the two years, with fatty acids profiles reflecting heterotrophic plankton production in 2007 and mostly autotrophic production in 2008. In agreement with Cushing's theory, our results highlight a match/mismatch, related to the food lipid quality rather than food quantity. For the first time, we show that the recruitment in marine bivalves may be dependent on phytoplanktonic pulses characterized by high levels of PUFA.     

Effects of salinity on the net uptake of zinc by the common mussel Mytilus edulis

The net uptake of zinc by the common mussel Mytilus edulis (L.) has been investigated under different natural and artificial salinity stresses. The effects of stable and fluctuating salinities on the uptake of zinc by the mussel are discussed in terms of three possible modes of action. Under certain highly-stressful conditions, salinity may affect the uptake of zinc by the mussel. This factor should be considered when the mussel is used as an indicator of environmental pollution by zinc in estuarine areas, or spurious conclusions may result.     

Ice nucleation in the blue mussel (Mytilus edulis)

The ice nucleation temperatures of different homogenised organs of the mussel Mytilus edulis L. were examined. The stomach and the hemolymph had the highest nucleation temperatures. In the homogenised stomach the nucleation temperatures were fairly constant throughout the year, whereas the nucleation temperatures of the hemolymph increased in the cold season. Bacterial growth experiments, microfiltration, and experiments using antibiotics indicated that the nucleators were not of bacterial origin. The nucleation temperatures of natural seawater were approximately −9 °C, whereas seawater made from distilled water and sea salt had nucleation temperatures of about −17 °C. The nucleation temperatures of the seawater were reduced when the seawater was filtered by the mussels. However, no clear indication that the nucleators in the stomach were obtained from the seawater was found. Stomach homogenates from mussels kept in nucleator-free water had the same supercooling points as stomach homogenates from mussels kept in natural seawater. This indicates that the nucleators in the stomach are not obtained from the seawater. The temperature and light conditions examined in the present study did not significantly influence the hemolymph ice nucleation temperatures of mussels kept in the laboratory. Received: 24 October 1996 / Accepted: 16 November 1996     

Simultaneous, long-term monitoring of valve and cardiac activity in the blue mussel Mytilus edulis exposed to copper

Valve and cardiac activity were simultaneously measured in the blue mussel (Mytilus edulis) in response to 10?d copper exposure. Valve movements, heart rates and heart-rate variability were obtained non-invasively using a Musselmonitor^® (valve activity) and a modified version of the Computer-Aided Physiological Monitoring system (CAPMON; cardiac activity). After 2?d exposure of mussels (4 individuals per treatment group) to a range of dissolved copper concentrations (0 to 12.5?M as CuCl₂) median valve positions (% open) and median heart rates (beats per minute) declined as a function of copper concentration. Heart-rate variability (coefficient of variation for interpulse durations) rose in a concentration-dependent manner. The 48?h EC₅₀ values (concentrations of copper causing 50% change) for valve positions, heart rates and heart-rate variability were 2.1, 0.8, and 0.06?M, respectively. Valve activity was weakly correlated with both heart rate (r?=?0.48?±?0.02) and heart-rate variability (r?=?0.32?±?0.06) for control individuals (0?M Cu²⁺). This resulted from a number of short enclosure events that did not coincide with a change in cardiac activity. Exposure of mussels to increasing copper concentrations (≥0.8?M) progressively reduced the correlation between valve activity and heart rates (r?=?0 for individuals dosed with ≥6.3?M Cu²⁺), while correlations between valve activity and heart-rate variability were unaffected. The poor correlations resulted from periods of valve flapping that were not mimicked by similar fluctuations in heart rate or heart-rate variability. The data suggest that the copper-induced bradycardia observed in mussels is not a consequence of prolonged valve closure.     

Relationship between lead concentrations in seawater and in the mussel Mytilus edulis: a water-quality criterion

Regression analysis on data collected from Port Phillip Bay and Western Port, Australia in 1979 shows that there is a significant equilibrium relationship between total recoverable lead in seawater and its concentrations in the mussel Mytilus edulis (P<0.001). The concentration of lead in seawater should not exceed 1.27 g l^-1 if the mussel is not to reach a lead concentration of 2.5 mg kg^-1 wet weight, a value frequently used as a food standard for human consumption. When a lead value of 2.5 mg kg^-1 wet weight is reached, the concentration factor by mussels for lead from seawater is 1969. The critical value of 1.27 g l^-1 could be used as a marine water-quality criterion for lead in waters where mussels are harvested.     

Oogenesis in the marine mussel Mytilus edulis: an ultrastructural study

Ultrastructural changes occurring during the course of development in oocytes of Mytilus edulis are described for mussels collected at monthly intervals over a period of one year (September 1981 to October 1982) from a site in Cornwall, England. During early stages of oogenesis the oocyte is surrounded by a small number of follicle cells but, as development proceeds, the follicle cells are restricted to the stalk region which attaches the oocyte to the acinar wall. Contact between the follicle cells and the developing oocyte is maintained by means of desmosomelike gap junctions. Organelles and inclusion bodies present in the ooplasm during oogenesis include rough endoplasmic reticulum (RER), Golgi bodies, mitochondria, free ribosomes, Balbiani's vitelline body, annulate lamellae and yolk and cortical granules. The RER, in particular, varies considerably throughout the course of development. Evidence for uptake of exogenous macromolecules into oocytes by pinocytosis is presented; it occurs in the basal region of previtellogenic oocytes prior to the formation of the vittelline coat. Lipid-yolk granules invariably have mitochondria in close association and, during the winter months, develop in close proximity to small, apparently glycogen-rich vesicles possibly suggesting that conversion of glycogen to lipid takes place in developing oocytes. Oocyte degeneration was commonly observed and involves initial breakdown of the plasma membrane followed by rupture of the vitelline coat. The oocyte contents once released into the acinar lumen are resorbed by the epithelial cells of the gonoducts, which are prevalent throughout the mantle of ripe individuals.