The combined effects of salinity and temperature on larvae of the mussel Mytilus edulis

The combined effects of salinity and temperature on survival and growth of larvae of the mussel Mytilus edulis (L.) were studied. The effects of salinity and temperature are significantly related only as the limits of tolerance of either factor are approached. Survival of larvae at salinities from 15 to 40 is uniformly good (70% or better) at temperatures from 5° to 20°C, but is reduced drastically at 25 °C, particularly at high (40) and low (20) salinities. Larval growth is rapid at a temperature of 15 °C in salinities from 25 to 35, at 20 °C in salinities from 20 to 35. Optimum growth occurs at 20 °C in salinities from 25 to 30. Growth decreases both at 25° and 10 °C; the decline is most drastic at high (40) and low (20) salinities.Part of a study completed at the Bureau of Commercial Fisheries, Biological Laboratory, Milford, Connecticut, USA, while on a UNESCO Fellowship.     

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.     

The effect of copper and zinc on the metabolism of the mussel Mytilus edulis

The effects of copper and zinc on the metabolism of the mussel Mytilus edulis (L.) and its component tissues were studied. 500 ppm copper sodium citrate inhibited oxygen consumption of the whole animal and gill tissue, but no similar effect was observed on digestive gland tissue. 500 ppm zinc sodium citrate exerted no effect upon gill or digestive gland respiration, and neither metal salt affected the respiration of homogenates of gill, digestive gland or gonad. Direct observation of gill tissues during exposure to the metals revealed that 500 ppm copper sodium citrate caused inhibition of ciliary activity; exposure of tissues to 2 ppm Cu for 24 h resulted in only partial inhibition of the cilía. It is suggested that metabolic suppression noted in whole animals and gill tissues is due to the inhibition of an energy-consuming process such as ciliary activity rather than interference with respiratory enzyme systems.     

Contribution to the ecotoxicological study of cadmium,copper and zinc in the mussel Mytilus edulis

A regulation of internal levels of some essential metals has been observed in various animals, whereas the bioaccumulation of several non-essential metals parallels their overloads in water. In the mussel Mytilus edulis L., we have attempted to determine if such a phenomenon exists by comparing the patterns of accumulation of copper and zinc vs cadmium. With this aim, mussels collected in the Bay of Bourgneuf (France) in November 1983 were exposed to these metals for 16 d. At external levels of zinc as high as 100 gl^-1, mussels were able to maintain a normal concentration in all groups of organs for 4 d. The ability of mussels to limit the bioaccumulation of copper and zinc varied from organ to organ, and decreased with higher levels of contamination and longer periods of exposure. In contrast, at the lowest experimental concentration and the lowest period of exposure, a significant increase of cadmium in mussel tissues was generally observed. Even at the highest sub-lethal doses, the levels of copper and zinc in mussel tissues were not much higher than the natural levels (contaminated:background ratios= 2.3 to 6.1), whereas the bioaccumulation of cadmium was less well restricted (contaminated:background ratios=136 to 192). The use of mussels as a bioindicator of pollution seems doubtful for essential metals, particularly as regards short-term pollution, since the levels of these trace elements in the organisms are largely independent of their concentration in the ambient seawater.     

Contribution to the ecotoxicological study of cadmium,lead, copper and zinc in the mussel Mytilus edulis

The understanding of natural fluctuations of metal concentrations in mussels used as bio-indicators is indispensable for a good assessment of the disturbances due to pollution. We have studied thoroughly the factors which condition bioaccumulation by using controlled populations of mussels, Mytilus edulis L., sampled monthly over more than two years (March 1982–May 1984) in the Bay of Bourgneuf, France. Seasonal changes in metal levels have been recognized, maximum values being observed in winter and early spring and minimum in later spring and summer. Depending on metal and size group, the ratios between these maximum and minimum values varied between 1.56 and 3.43. The fluctuations in soft-tissue weight appear to be the main explanatory factor of seasonal variations in metal concentrations in mussels. Fluctuations in metal levels related to size of mussels were observed. Except for cadmium in mussels with a mean dry weight of soft tissues >0.2 g, a slight decrease in metal concentrations was observed for growing individuals: the regression coefficient b was -0.10, -0.11, -0.13 and -0.27 for Cd, Cu, Zn and Pb, respectively. The metals examined were rather uniformly distributed among the different groups of organs (visceral mass, gills and palps, remainder) and, consequently, several analyses of metals in different organs do not provide much more information than one analysis on the whole soft tissues. It is concluded that fluctuations related to size or season are reflected by only moderate differences in the maximum and minimum concentrations of metals in the mussels, but that they are nevertheless sufficient to conceal low chronic or short-term pollution, except at those sites where the normal environmental conditions are well-documented.     

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     

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.     

Synergistic effects of lead,salinity and temperature on embryonic development of the mussel Mytilus galloprovincialis

Combined effects of lead, salinity and temperature on the embryonic development of the mussel Mytilus galloprovincialis Lmk. were studied under laboratory conditions. The basic experimental design was a 4x6 factorial experiment using 4 lead concentrations (100, 250, 500 and 1000 ppb Pb²⁺) and 6 salinity levels (from 25 to 37.5 with 2.5 intervals). These factorial designs were carried out at three constant temperatures (150, 17.50 and 20°C). The statistical analysis indicated that salinity changes have more effect on the embryonic development than temperature. Optimal development was observed at 34.8 and 15.6°C, which is in accordance with observations in the field. The effect of lead was mininal in optimal salinity and temperature conditions. The deleterious effect of lead on the embryonic development was especially conspicuous at 20°C. Since in nature spawning occurs at temperatures inferior to 20°C, lead will probably not drastically decrease the potential recruitment of mussel spat in the littoral populations of the northern Adriatic Sea, where the salinity of the water is relatively stable. Under experimental conditions, lead caused a delay or inhibition of the embryonic development with the occurrence of a large number of abnormal larvae.     

The reproductive cycle and physiological ecology of the mussel Mytilus edulis in a subarctic,non-estuarine environment

Mytilus edulis L. occupies a habitat in eastern Newfoundland, Canada, which is characterised by low temperatures and a lower concentration of particles in suspension than is found in estuarine locations. Gametogenesis occurs in the spring, and energy reserves from the previous year are not utilised in the synthesis of gametes. Oxygen uptake is not elevated during the winter, in contrast to most European populations, which undergo gamete development at that time. There is little seasonal variation in clearance rate. This may be attributable in part to the absence of the high particulate load which has been shown to depress feeding activity in some estuarine populations. Energy balance is positive at all times of the year, and conversion efficiency is high, suggesting that conditions are favourable. A number of physiological comparisons are made between Newfoundland mussels and those from European waters.     

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.     

The common mussel Mytilus edulis as an indicator of pollution by zinc,cadmium, lead and copper. II. Relationship of metals in the mussel to those discharged by industry

Concentrations of zinc, cadmium, lead and copper were determined in whole soft parts of the common mussel Mytilus edulis (L.) sampled in Port Phillip Bay and Western Port Bay, both in Victoria, Australia. The mussels were sampled according to procedures suggested by previous studies in order to eliminate the effects of natural environmental variables. Results of the analytical studies were compared to data on the quantities of trace metals known to be discharged by industry into the catchments of each Bay. This allowed an evaluation of the indicator ability of the mussel without the need for multiple analyses of water samples. The results suggest that the mussel is capable of acting as an efficient time-integrated indicator of zinc, cadmium and lead over a wide variety of environmental conditions. Although additional variables may remain to be investigated, the mussel is recommended as an alternative to the analysis of water and sediments. With further development this organism may also be useful for monitoring industrial effluent characteristics at the discharge site, allowing rapid, reliable and inexpensive control of water quality. In contrast, the mussel is of doubtful use as an indicator of copper; other indicator types such as macroalgae may be better-suited to the study of this element.     

Aspects of nitrogen metabolism of the common mussel Mytilus edulis: Adaptation to abrupt and fluctuating changes in salinity

Mytilus edulis L. were exposed to abrupt (3015 and 1530) and fluctuating (sinusoidal 12 h cycles of 301530) changes in salinity, and the changes in the total osmoconcentration of the haemolymph were recorded. The response of nitrogen metabolism to the altered extracellular osmotic concentrations was investigated in terms of the concentrations of the total NPS (ninhydrin-positive substances) pool and the individual amino acids of the tissues, the concentration of the amino acids of the haemolymph, and the rates of excretion of ammonia and amino acids by whole individuals. The haemolymph became isosmotic with the seawater with abrupt changes in salinity, but with fluctuating salinity was slightly hyperosmotic as the salinity decreased and then slightly hypo-osmotic as the salinity increased. This resulted in a reduction in the extent of the extracellular osmotic change compared to the change in fluctuating salinity to which it was exposed. Total NPS of the tissues decreased with an abrupt decrease in salinity and increased with an abrupt increase in salinity, but a seasonal dependence of the response was indicated. The short-term response of tissue NPS to fluctuating salinity was equivocal, but with long-term exposure the concentration declined. Ammonia and amino acid excretion increased with both an abrupt decrease in salinity and fluctuating salinity and decreased with an abrupt increase in salinity. Haemolymph amino acids increased with an abrupt decrease in salinity. The increased rates of nitrogen excretion accounted for the reductions in the NPS concentrations of the tissues except in the early stages of fluctuating salinity. Taurine, aspartate, threonine, serine, glycine and arginine declined with an abrupt decrease in salinity while alanine and glutamate increased slightly. With an abrupt increase in salinity, alanine and ammonia accumulated in the tissues and then declined while the other amino acids increased slowly over a longer time-course. Similar individual amino acid responses were seen with long-term exposure to fluctuating salinity, except for taurine which did not decrease in concentration. On the basis of the changes in tissue amino acids and ammonia, it is suggested that the alanine dehydrogenase reaction is the primary nitrogen-fixing reaction in marine bivalves such as M. edulis.     

Uptake of cadmium and zinc by the mussel Mytilus edulis and inhibition by calcium channel and metabolic blockers

The mussel Mytilus edulis is extensively used to monitor metal contamination of estuarine and marine systems. Nonetheless, the mechanisms of metal uptake are poorly understood. To characterise the systems involved in cadmium and zinc uptake, the interaction between the two metals and the effects of different calcium channel blockers (diltiazem, nifedipine, verapamil) and inhibitors of active transport and metabolism (ouabain, sodium cyanide, 2,4 dinitrophenol) on the uptake of calcium, cadmium and zinc in Mytilus edulis have been studied. To separate direct from indirect effects of the inhibitors on metal uptake, their influence on the physiological condition of the mussels was also investigated. This was done by measuring clearance, respiration and excretion rates under the different exposure regimes and determining the scope for growth as an integrative index for physiological condition. The study has shown that the uptake of cadmium and zinc by Mytilus edulis can be modulated by calcium channel blockers and other inhibitors. The inhibitors also influenced physiological condition, but a significant correlation with the effects on metal uptake did not exist in most cases. Cadmium and zinc also inhibit each other's uptake, but the type of inhibition is not yet clear. The effects of the inhibitors on cadmium and zinc uptake are very different from the effects on calcium uptake, indicating that cadmium and zinc are preferentially taken up through other gateways. Overall, a significant degree of linear association is found between the effects of the inhibitors on the uptake of cadmium and zinc, suggesting common uptake routes. In addition, the effects of the calcium channel antagonist on the uptake of the metal ions are organ dependent, indicating that other types of channels are involved in the uptake of the metal ions in the gills and digestive system. Received: 16 February 1999 / Accepted: 23 September 1999     

Growth,condition and specific dynamic action in the mussel Mytilus edulis recovering from starvation

Juvenile Mytilus edulis were grown individually in plastic racks in a tidal salt marsh for 72 d in 1984, starved in the laboratory for 130 d, and then fed the alga Isochrysis galbana daily for 64 d. Oxygen consumption was measured at various times during the course of starvation and recovery. The effects of both size (tissue dry weight) and condition (tissue size relative to shell size) on the rate of oxygen consumption during the course of starvation and recovery were analyzed by multiple regression. Weight-specific preprandial rate was inversely correlated with both size and condition. Weight-specific active rate (measured shortly after feeding) was correlated with size but not condition. Relative Scope for Activity was inversely correlated with size and positively correlated with condition. Relative Specific Dynamic Action (RSDA; the integrated physiological and mechanical response to a meal) was initially correlated negatively with size and subsequently positively with condition. Glycogen content was shown to be positively correlated with condition in mussels before starvation and during recovery. During recovery, experimental mussels returned to 90% of their estimated dry weight prior to starvation, and from 53% after starvation. At weekly intervals during recovery, oxygen consumption was measured following a meal until it returned to the preprandial rate. Both pre- and postprandial volumes (l at STP) of oxygen consumed per hour ( O _2pre and O _2post , respectively) increased significantly during recovery. The postprandial rise in oxygen consumption increased significantly from 15% to 23% of O _2pre . At the end of the recovery period, RSDA [( O _2post – O _2pre )/ O _2pre ] was independent of final dry weight, but was significantly correlated with percent dry weight recovered (r ²=0.44; df=10; P<0.02). The increase in RSDA may reflect increased utilization of food and its conversion to soma during recovery from starvation, as distinct from mechanical energy expenditure (feeding activity) following a meal.     

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.     

Relationships between seston,available food and feeding activity in the common mussel Mytilus edulis

The feeding and metabolic rates of Mytilus edulis L. of different body sizes were measured in response to changes in particle concentrations ranging from 2 to 350 mg l^-1. Rates of oxygen consumption were not significantly affected by changes in seston concentration, whereas clearance rates gradually declined with increasing particle concentration. Pseudofaeces production was initiated at relatively low seston concentrations (<5 mg l^-1). Marked seasonal changes were recorded in the composition of suspended particulates (seston) in an estuary in south-west England. Total seston was sampled at frequent intervals throughout an annual cycle and analysed in terms of: particle size-frequency distributions, total dry weight (mg l^-1), inorganic content, chlorophyll a, carbohydrate, protein and lipid. The particulate carbohydrate, protein and lipid content provided an estimate of the food content of the seston. The results are discussed in terms of the food available to a nonselective suspension feeder, such as M. edulis, during a seasonal cycle. The effect of inorganic silt in suspension was mainly to limit by dilution the amount of food material ingested rather than to reduce the amount of material filtered by the mussel. In winter, the food content of the material ingested was 5%, and this increased to 25% during the spring and summer.     

Ultrastructural changes in the lysosomal-vacuolar system in digestive cells of Mytilus edulis as a response to increased salinity

The effects of increasing salinity on the ultrastructural morphology of the lysosomal-vacuolar system in digestive cells of the common mussel Mytilus edulis were investigated in order to relate structural changes to previous biochemical and cytochemical observations. After 3 h of increased salinity, from 21 to 35%., the digestive cells showed an increase in numbers of heterolysosomes. There was some evidence of digestive cell breakdown, the contents forming membrane-bound bodies and being released into the tubule lumen. After 12 h of increased salinity, heterolysosomes were prevalent in the digestive cells. There was increased evidence for digestive-cell breakdown, many of the tubule lumina being packed with membrane-bound bodies. It is concluded that increasing salinity from 21 to 35%. stimulates the lysosomal-vacuolar system, as a result of autophagocytosis or apoptosis; this is consistent with the hypothesis that intracellular, lysosomally-mediated, catabolism of proteins is a source for free amino acids during the adaptation of mussels to increased salinity.     

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.     

Byssus drifting and the drifting threads of the young post-larval mussel Mytilus edulis

The long, drifting threads secreted by young post-larval mussels (Mytilus edulis L.) are simple monofilaments, distinct in form and function from the attachment byssus threads. The diameters of both thread types are in the micron or sub-micron range but, whereas the attachment threads are of restricted length and terminate in an attachment plaque, drifting threads exceed the post-larva in length by more than two orders in magnitude and are without plaques or any other structures. Transmission electron micrographs of drifting threads show no evidence of internal sub-structure. In contrast, attachment threads appear to be made up of filaments. These studies confirm that the drifting threads are highly effective in enhancing the dispersal of young mussels. The terminal sinking velocity of young drifters is typically ca. 1 mm s^-1. At this velocity the suspension range above the sea bed, assuming given values of vertical diffusivity, is estimated to be 0.5 to 5 m. Calculations of the fluid drag experienced by post-larvae in the water column show that the theoretical viscous drag force on the thread is sufficient to account for the reduced sinking rate of drifters. The calculated contribution of the thread to the total drag is approximately one order of magnitude greater than that of the post-larval body. A rapid thread-deployment strategy, shown by post-larvae which are brought into suspension, may prolong each drifting excursion and thus further enhance dispersal in turbulent marine environments.