Feeding and cleaning mechanisms in the suspension feeding bivalve Mytilus edulis

Feeding and pseudofeces formation were studied in intact Mytilus edulis clearing suspensions of graphite particles, and the processes were compared with activities observed in mussels with a severed adductor muscle. Most particulate material in the stomach was present in the suspended state. Ingestion of particles in suspension could take place concurrently with the production of pseudofeces. Severing of the adductor muscle resulted in profuse secretion of mucus that gradually subsided to a constant low level. Addition of graphite particles at concentrations that did not cause mucus secretion in the intact mussels strongly stimulated secretion in the operated mussels, resulting in the accumulation of highly viscid, mucusparticle aggregates. These aggregates were mechanically stable, in contrast to the fragile pseudofeces. It is indicated that normal feeding depends upon hydromechanical mechanisms that produce highly concentrated suspensions of particles for ingestion, and that mucociliary mechanisms serve to clean the gills and other organs of the mantle cavity for excess particulate material. It is further indicated that intact mussels secrete mucus only in the amounts needed to consolidate excess particulate material, and that lesions affect the normal balance between particles in suspension and mucus secretion by enhancing the sensitivity of the mechanisms that control mucus secretion.     

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.     

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.     

Introgression and mitochondrial DNA heteroplasmy in the Baltic populations of mussels Mytilus trossulus and M. edulis

A strong clinal change in salinity occurs between the Baltic Sea and the North Sea, Atlantic Ocean, in the Danish Straits, where hybridization zone between mussels Mytilus edulis and M. trossulus has been reported. Eleven samples of mussels were studied from the Danish Straits and the inner Baltic Sea. Extensive introgression of M. edulis alleles from the North Sea into populations throughout the Baltic was ascertained for mitochondrial DNA (mtDNA) and two nuclear markers (ME15–16 and ITS). In the opposite direction, introgression of M. trossulus alleles into the M. edulis background was observed at the EFbis nuclear marker in populations from Kattegat (Danish Straits). While only M. edulis F (female) mtDNA was present in the Baltic, there were still strong differences in frequencies in the control region length variants between the Danish Straits and the inner Baltic samples, and weaker variation in coding region ND2–COIII haplotype frequencies. In the assays of the two mtDNA regions, various patterns of heteroplasmy were detected in 32% of all the studied individual mussels; this includes the presence of distinct, independently inherited M and F mitochondria in males, as well as the presence of two different distinguishable F genomes. The male-inherited M mtDNA genomes are quite common in the mussels from the Danish Straits, but very rare in males from the inner Baltic. Instead, a recombined control region variant (1r), which seems to have taken over the role of the M genome, was present in a number of specimens in the Baltic. Observations of heteroplasmy for two F genomes in some females and males confirm disruptions of the doubly uniparental inheritance mechanism in the hybrid Baltic Mytilus.     

A proteomic approach to the study of the marine mussels Mytilus edulis and M. galloprovincialis

In this study, a proteomic approach was applied for the generation of reference maps and subsequently to detect, quantify and compare the global protein expression between two related species of marine mussels, Mytilus edulis and Mytilus galloprovincialis, growing in their own geographical habitats. A comparative study of the protein profiles generated from analytical two-dimensional electrophoresis gels was performed, and changes in protein expression were analysed quantitatively by computer analysis. An average of 1,278 spots per gel was detected in 16 individuals (8 M. edulis and 8 M. galloprovincialis); however, not all spots were included in the study. Expression of 420 spots was compared, and significant differences in the intensity levels were detected in 37 protein spots (8.8%). Fifteen proteins showed higher expression in M. edulis, and 22 proteins, in M. galloprovincialis. The technique of peptide mass fingerprinting using MALDI-TOF (matrix-assisted laser desorption ionisation/time-of-flight) and/or nanoelectrospray double subfragmentation mass spectrometry enabled the unambiguous identification of 15 of these 37 differentially expressed proteins. Most of the identified proteins can be grouped basically into four broad functional classes: cytoskeletal and myofibrillar proteins, proteins associated with stress response, proteins associated with the storage or production of energy, and proteins related to rearrangement in the synthesis of native structures. These results expand our understanding of the molecular differentiation of the two mussel taxa and serve as a useful base for future ecological, physiological and genetic studies.     

Influence of algal concentration and temperature on the filtration rate of Mytilus edulis

The filtration rates of Mytilus edilis (=galloprovincialis; 40 mm) were determined in relation to food concentration and temperature, using pure suspensions of the unicellular alga Platymonas suecica in concentrations ranging from 3x10⁵ cells/l to 1.5x10⁸ cells/l. The rate of filtration (ml/h/mussel) generally decreased as cell concentrations increased, and dropped to low values when concentrations above 5x10⁷ cells/l were supplied. The amount of water swept clear varied continuously, and noticeable differences in the filtration activity of M. edulis were observed over short time intervals (5 min). Fluctuations of filtered volumes per unit time were greater with lower than with higher concentrations of algae. The influence of temperature on filtration activity was highest between 5°–15°C and 25°–30°C. A temperature increase from 15° to 25°C resulted in only a slight increase in filtration rate. At 5° and 30°C, filtration dropped to very low values, namely 350 and 100 ml/h, respectively. The temperature coefficients for the filtration rates of M. edulis were determined as: Q₁₀ (5° to 15°C)=4.96; Q₁₀ (10° to 20°C)=1.22. The amount of algae cells ingested per mussel per hour is directly related to food concentration. The maximum number of cells filtered/mussel/h in an algal suspension of 70x10⁶ cells/l was 21.5x10⁵ cells/h. Cell concentrations of up to 40x10⁶ cells/l were swept clear without producing pseudofaeces. The critical cell density for M. edulis was reached at algal concentrations of 70 to 80x10⁶ cells/l. Above these concentrations no normal filtration activity was observed.     

The effect of chelating agents on the uptake and accumulation of cadmium by Mytilus edulis

The uptake, storage and excretion of cadmium by the common mussel Mytilus edulis (L.) has been studied at sub-lethal concentrations using the radioactive isotope ^115mCd as a marker. After an initial lag period, the uptake at low concentrations in sea water is linear with time and directly proportional to the sea water concentration, with a maximum concentration factor of 165 at 0.7 g Cd/ml sea water. A decrease occurs at higher concentrations indicating saturation of the available binding capacity. Prior complexation of the cadmium with either EDTA, humic and alginic acids or pectin doubles both the rate of accumulation and the final tissue concentrations (order: kidneyviscera>gillsmantle>muscle) and eliminates the lag period, suggesting that ionic cadmium must first be complexed before uptake can occur. A mechanism for this effect, which may involve thionein, is described. The rate of excretion of cadmium is 18 times slower than that of uptake, with the major route via the kidney but not via the byssal threads as with particulate iron. The need to detoxify and store cadmium by an immobilization mechanism is a consequence of this slower rate of elimination.     

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.     

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.     

Effects of temperature,nutritive and metal stressors on the reproductive biology of Mytilus edulis

Experimental observations suggest that the gametogenic cycle of Mytilus edulis from Swansea Bay, South Wales (U.K.) is adversely modified by the combined effects of temperature, nutritive and sublethal metal-induced stress. During low temperature stress, oogenesis remained in the vegetative stage and vitellogenesis only commenced when temperatures were raised to ambient levels. Copper was the most toxic metal, although its uptake was slowest. A concentration of 0.05 mg l^-1 of this metal proved lethal at ambient temperatures, although at low temperatures there was no mortality. Copper suppressed both the growth of young oocytes and vitellogenesis in larger oocytes. Zinc was less toxic than copper, although this also inhibited oocyte development and resulted in severe lysis of gametes. Cadmium was the least toxic of the three metals studied and suppressed gametogenesis only in the initial stages of gonad development. The low toxicity of cadmium to gametogenesis may be a result of its high solubility and its possible storage in the digestive gland.     

Role of the kidney in determining the whole soft tissue zinc concentration of individual mussels (Mytilus edulis)

Mussels (Mytilus edulis) were collected from an unpolluted site in Newfoundland in 1984 and 1985 in order to determine the zinc levels of the whole soft tissues and individual organs. Individuals were collected in such a way as to eliminate all known physiological and environmental factors which might have led to differences in zinc concentration between individual mussels. The inherent variability of the whole soft tissue zinc concentration was found to be very high with a coefficient of variation (CV) around 40%. Most individual organs had relatively low mean zinc concentrations with small values for CV (10–18%). However, the kidney showed a relatively high mean zinc concentration (828 ppm) with a remarkable degree of variability (94–3 410 ppm) (CV=78%) and is responsible not only for the high degree of inherent variability in the population but also for the high levels of zinc found in the whole soft tissue of some individual mussels. Those individuals with high whole soft tissue zinc concentrations often had high kidney zinc concentrations and stored as much as 53% of their zinc load in the kidney (which accounted for only 1–3% of the total weight). However, other individual mussels with low whole soft tissue zinc concentrations had low kidney zinc concentrations and stored as little as 3% of their body zinc load in the kidney. Hence, the role that the kidney plays in zinc storage varies markedly from mussel to mussel.M.S.R.L. Contribution No. 624     

The occurrence and effects of Mytilicola intestinalis in Mytilus edulis

Mytilicola intestinalis was observed in the mussel Mytilus edulis in increasing numbers for the first time at Brighton (England), in October 1966; the populations here and at Whitstable were examined. Mussels exposed high in the littoral zone were less heavily infected than those lower down, the degree of infection being directly related to the duration of exposure in each tidal cycle. Silt in the intestine of the mussel is considered to act as a controlling factor in numbers of parasites present at Whitstable. Egg-bearing copepods were present in samples throughout the year, suggesting that breeding is not interrupted by the winter. Evidence indicates that juvenile stages of the parasite cause most damage to the host, due in part to their presence in the ramifications of the hepatopancreas. Recovery of the mussel from the effects of parasitation is rapid, following a reduction in parasite population density and number of juveniles. In the laboratory, M. edulis is more rapidly affected by lack of food at 10 °C than M. intestinalis. No dead parasites were seen during 4 months of laboratory storage. Juvenile parasites continned to mature, indicating that this period of time may be required for Mytilicola intestinalis to reach maturity at 10°C.     

Distribution of Mytilus edulis, M. galloprovincialis, and their hybrids in open-coast populations of mussels in southwestern England

The distribution of Mytilus edulis Linnaeus, M. galloprovincialis Lamarck, and their hybrids was examined in mussel populations in southwest England in 1996 and 1998. This is a region where both parental taxa and populations containing large numbers of hybrids co-occur yet a fine-scale mapping of the hybrid populations has not been conducted. In this study the geographic distribution of hybrid populations was determined in southwest England over 360 km of coast from Tintagel Castle in north Cornwall to Beer in south Devon. Sample localities were spaced from 5 to 20 km apart. Genotypes for individual mussels were determined using PCR to amplify a size polymorphism at the Glu-5' locus that is completely differentiated between M. edulis and M. galloprovincialis. Hybrid populations, characterized by high frequencies of individuals with heterozygous genotypes and a pattern of decreasing frequency of M. edulis alleles with increasing shell length, were continuously distributed along 180 km of open coast in southwest England. This "hybrid patch" was bordered at one end by geographically extensive populations of pure M. edulis and at the other end by nearly pure populations of M. galloprovincialis. Strong natural selection in hybrid populations results in a decline in the frequency of M. edulis alleles with increasing size. Wave-exposure has previously been implicated as the agent producing this pattern of selection, but in the present study the relationship between allele frequency and body size was not correlated with variation in wave shock intensity among localities within the hybrid zone. The transition between hybrid populations and those containing pure populations of M. edulis or M. galloprovincialis is abrupt which suggests that coastal circulation patterns may provide strong barriers to larval dispersal which accounts for the position and maintenance of the hybrid zone.     

Geographic variation,niche breadth and genetic differentiation at different geographic scales in the mussels Mytilus californianus and M. edulis

Genetic differentiation was investigated in the marine mussels Mytilus californianus Conrad and M. edulis Linn. from the west coast of North America. In allopatry with M. californianus, M. edulis occurs throughout the intertidal zone; however, in microgeographic sympatry its ecological range is restricted to above the M. californianus mussel bed and to patches of substratum opened by natural disturbances within the bed. Over the same geographic scale, the broader-niched M. edulis shows greater among-locality genetic difference and greater levels of polymorphism than M. californianus at two enzyme loci. Genetic differentiation on a geographic basis was investigated in M. californianus at a single rock (on a scale of meters), on an island (on a scale of kilometers), throughout a strait (on a scale of 10² kilometers), and along the west coast of North America (on a scale of 10³ km). Differentiation was minimal over the west coast, and could be explained by microhabitat differences in a local area. The minimal differentiation in west coast mussels relative to strong geographic differentiation of M. edulis on the east coast of North America may be related to the steeper latitudinal thermal gradient on the east coast. Local ecologically related microgeographic variation can result in biased and misleading estimates of genetic heterogeneity but microgeographic variation at enzyme loci may not be due to selection at the loci investigated or may even be due to the breeding structure of the mussel populations, as suggested by Tracey et al. (1975).     

Effect of temperature and food on the heart beat,ventilation rate and oxygen uptake of Mytilus edulis

The heart beat, ventilation rate and oxygen uptake of Mytilus edulis L. were measured simultaneously, in response to changes in temperature and food level. There was no thermal acclimation of heart-beat frequency or amplitude to temperatures from 5° to 25°C. Oxygen consumption and ventilation rate acclimated to 10°, 15° and 20°C, but not to 25°C. Starvation reduced the rate of oxygen uptake and heart-beat frequency to a standard level and, in response to food, the ventilation rate and oxygen consumption immediately increased to an active level. Feeding was maintained after the initiation of active metabolism, and during the following 10 days the heart-beat frequency gradually increased to the level characteristic of fed individuals. There was no direct correlation between the rate of oxygen consumption and heart rate, and an apparent absence of a close nervous coupling between ventilation rate and heart rate in M. edulis.     

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.     

Very long-chain aliphatic hydrocarbons in lipids of mussels (Mytilus edulis) suspended in the water column near petroleum operations off Sable Island,Nova Scotia,Canada

Mussels (Mytilus edulis) suspended in the water column in 1994 and 1995 for the monitoring of oil drilling operations off Sable Island, Nova Scotia were examined for hydrocarbon profiles, particularly aliphatic hydrocarbons. A spring bloom of phytoplankton occurred during the 90-d suspension period in 1995. Hydrocarbons isolated from the 1995 suspended mussels showed very high concentrations of both biogenic hydrocarbons and very long-chain n-alkanes from C₂₀ to C₃₂, initially thought to be petrogenic. Both types of hydrocarbons were either not detected or were only present in trace amounts in the mussels suspended in 1994 at similar sites. The biogenic hydrocarbons in the 1995 mussels were apparently of planktonic origin, from the spring bloom, and were dominated by heneicosahexaene (21:6), followed by pristance, heptadecane, and varions monounsaturated and polyunsaturated phytenes, heptadecenes, nonadecenes and heneicosenes. They could be readily hydrogenated to yield the basic alkanes. The 1995 mussels suspended within 1 km from the oil well platform were probably slightly tainted by petrogenic hydrocarbons, as evidenced by the detection of phytane and high concentrations of total aliphatic hydrocarbons, whereas the mussels suspended 10 km from the platform showed only high concentrations of biogenic hydrocarbons and the novel long-chain n-alkanes. The occurrence of an unusual phytoplankton bloom during the suspension period severely interfered with the petroleum monitoring role of mussels by altering the mussel hydrocrbon profiles through the accumulation into and probably selective depuration of xenobiotic hydrocarbons from the mussel, tissues.J. Parsons (deceased)     

The significance of temperature,salinity and zinc as lethal factors for the mussel Mytilus edulis in a polluted estuary

Mussels, Mytilus edulis L., were subjected to high temperatures, low salinities and dissolved zinc in order to investigate possible environmental hazards of a discharge of heated effluent near Newport on the Yarra River estuary, Victoria, Australia. Exposure to zinc at 0.8 mg l^-1 for 14 d in otherwise favourable conditions significantly increased mortality resulting from subsequent exposure to temperatures between 29° to 31°C for 24 h without added zinc. Mussels collected from water of temporarily lowered salinity (8–16 S) showed significantly lower thermal resistance than controls collected from marine salinities (35 S). Mussels taken from a marine environment and exposed to 10 S died at a rate which increased with temperature. Mussels acclimated for 14 d to combinations of 10°, 16° and 22°C and 22 and 35 S, and subsequently exposed to increased zinc concentrations accumulated zinc to levels which were independent of temperature and salinity. The zinc was lethal more quickly at 22°C and 35 S than at the lower temperatures and salinities. The modes of toxic action of the salinity, zinc and temperature factors are discussed and it is argued that zinc which has been found accumulated in mussels near Newport could be reducing their resistance to raised temperatures and perhaps other stresses, probably as a result of effects on lysosomal functioning. The evidence suggests that the heated effluent will accelerate any toxic effects of zinc or low salinities which occur near Newport and so poses a hazard in winter as well as in summer.