Bioseston production by Mytilus edulis and its effect in experimental systems

Mytilus edulis L. (Bivalvia: Mollusca) can produce appreciable quantities of 3.4 to 5.4 m diameter bioseston, when previously fed on Phaeodactylum tricornutum and Tetraselmis suecica. Bioseston production, observed in flowing filtered sea water, is shown to originate from resuspension of faeces and can cause filtration-rate underestimates of up to 35% under certain experimental conditions. Feeding with Thalassiosira pseudonana below a concentration of 4x10⁷ particles l^-1 caused no significant bioseston production and this alga would, therefore, be suitable for use in quantitative estimates of filtration rate.     

Degradation of bacteria by Mytilus edulis

The uptake of several species of bacteria by the common mussel Mytilus edulis (L.) and the subsequent fate of some polymers of the bacteria have been investigated in a study carried out during 1981. Bacteria (Escherichia coli, Micrococcus luteus, M. roseus, Bacillus cereus, Staphylococcus aureus and a marine pseudomonad, 1-1-1) were radiolabelled by growth in medium containing ³H-thymidine and the uptake of bacteria by Mytilus edulis was monitored. Labelled and unlabelled bacteria, at initial concentrations of 0.5 to 1x10⁷ bacteria ml^-1, were cleared at similar, exponential rates with no significant difference in the rates for different bacteria: 90% of bacteria were cleared in a mean time of 1.93±0.12 h (SEM, n=63). Those bacteria with cell walls which were sensitive to M. edulis lysozyme were rapidly degraded by the mussel and ³H-labelled DNA was released in a form not precipitable by 10% trichloroacetic acid. Lysozyme-resistant bacteria (Micrococcus roseus and S. aureus) were cleared from suspension by Mytilus edulis but most were rejected intact. By measuring the rate of release of ³H-thymidine-labelled material from the mussel the rate of degradation of lysozyme-sensitive bacteria by M. edulis was found. For different bacteria the degradation rate varied from approx 2x10⁸ to 27x10⁸ bacteria h^-1 with an overall mean of 10x10⁸ bacteria h^-1. A thymidine- and diaminopimelicacid-requiring auxotroph of E. coli was radiolabelled with ³H-thymidine, ³H-diaminopimelic acid or ¹⁴C-glucose and fed to M. edulis. Bacteria were cleared and degraded by the mussel; ³H-diaminopimelic acid-labelled or ¹⁴C-glucose-labelled polymers were retained, whereas ³H-thymidine-labelled polymers were released into the surrounding water. Extracts of the digestive gland of M. edulis degraded lysozyme-sensitive bacteria to release ³H-thymidine-labelled material, but did not release ³H-thymidine-labelled material from lysozyme-resistant bacteria. It is concluded that M. edulis can select lysozymesensitive bacteria for subsequent processing and discriminate between bacterial polymers to reject DNA. Also, bacteria could provide a substantial fraction of the carbon requirement of the mussel.     

Morphological variation of Mytilus edulis and Mytilus trossulus in eastern Newfoundland

Allopatric populations of Mytilus species show distinct shell morphology which may be due to genetic and/or environmental effects. Sympatric populations of Mytilus species show similar shell morphology which may be due to hybridization eroding morphological differences and/or the influence of common environmental conditions. The present study examined shell morphology and shell shape from 16 sites in eastern Newfoundland where M. edulis L. and M. trossulus Gould coexist in common environments with limited hybridization. Shell morphology was based on measurements of eight characters, and shell shape was quantified by elliptic Fourier analysis of shell outlines. Significant differences were observed between species for both shell morphology and shell shape across 16 sites sampled. The relatively small differences in morphology and shape between the species were probably due to exposure to common environments rather than hybridization. Shell shape for M. edulis was more eccentric compared to M. trossulus which was more elongated. Shell shape analysis of a range of size classes at one site showed a change from an eccentric to an elongated shape going from the smaller to the larger size classes. Both species showed a similar trend, with the larger M. edulis more eccentric and the larger M. trossulus more elongated. Received: 17 July 1998 / Accepted: 6 January 1999     

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.     

Temporal patterns of shell-gape in Mytilus edulis

Hitherto published evidence of the presence or absence of endogenous activity rhythms in bivalve molluscs is equivocal. Mytilus edulis L. were collected from a North Wales (UK) estuary in 1985, and shell-gaping was investigated in individual mussels under constant conditions in the laboratory. Results suggest that there is no endogenous circatidal rhythm of shell-gaping in this species, This is consistent with the view that, unlike mobile species, sessile intertidal species are much more likely to exhibit exogenous rather than endogenous responses to tidal fluctuations. There is some evidence of weak circadian rhythmicity of shell-gaping in M. edulis, with greater duration of shell-closure during hours of expected daylight. Such behaviour could represent an adaptational defence against visually-feeding predators.     

Uptake and cellular distribution of cadmium in Mytilus edulis

Cadmium uptake has been studied in starved and fed Mytilus edulis L. It is suggested that fairly elevated cadmium contents in fed mussels are not due to contaminated food, but to increased pumping rate when food is available. Highest concentration and main body burden are found in the mid-gut gland. Transport via haemolymph, and selective discrimination at the basement lamina of the mid-gut gland tubuli are regarded as mainly responsible for accumulation. Mercury seems to be processed in a similar way as cadmium. In the tubuli, both metals are immobilized in membrane-bound vesicles, which are finally defaecated.This work was financially supported by the German Research Council (DFG) (Sto 75/4 and Th 158/13).     

Rate of heat dissipation by gametes and larval stages of Mytilus edulis

Heat dissipation of sperm, eggs and various stages of the planktonic larvae of the mussel Mytilus edulis L. was recorded directly at 15°C. The absolute heat dissipation increased steadily over all stages examined. The massspecific heat dissipation also increased during early embryogenesis from the fertilized egg to the D-larval stage. Starting from the early veliconcha stage, however, the mass-specific rate of heat dissipation declined from about 90 m W g^-1 with increasing body size with a weight exponent of-0.35. A comparison with literature data revealed a consistently higher metabolic activity than previously estimated by indirect calorimetry for gametes and larvae.     

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.     

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.     

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.     

An electron-microscope study of periostracum repair in Mytilus edulis

The ultrastructural details of changes that occur in the inner face of the outer fold of Mytilus edulis during periostracum repair are described. Initially, after the periostracum is slit, increased secretory activity occurs in the cells, apparently controlled by lysosomes. Proto-ostracal material deposited outside the cells seems to be due to Golgi secretory granules, which dissociate and then re-assemble outside the cells to form an organised layer. During the later stages of repair, autophagic vacuole formation increases in the cells, possibly because of increased demands for synthetic material. When substantial amounts of proto-ostracal material are secreted and the outer fold is sealed off from the external environment, the cells return to their normal condition. During the early stages of repair, amoebocytes appear to be implicated in the laying down of proto-ostracal material.     

Growth and size structure in a baltic Mytilus edulis population

Since Mytilus edulis L. has very few predators and competitors for space, it has become a biomass dominant in the Baltic proper covering hard substrates from the water surface to more than 30 m depth. In order to investigate the factors controlling size and production in a Baltic M. edulis population, growth was studied by the analysis of annual growth rings, measurements of caged individuals and the analysis of size classes in the population, and on settlement ropes. The total number of mussels in a representative mussel bed at 4 m depth varied between 36 000 and 158 000 ind · m^-2 during the year, mainly due to variations in very small mussels (<2 mm), whereas the abundance of mussels 2mm was rather constant between about 17 000 and 28 000 ind · m^-2. Maximum numbes of mussels < 2 mm, amounting to 132 000 ind · m^-2, were found after settlement in summer, but still half a year later in spring, 65 000 ind · m^-2 < 2 mm were registered, due to very strong intraspecific competition for food and space leading to the competitive suppression of small individuals and large variations in growth rates. Due to the special size-structure of the population only the analysis of annual growth rings could be used to estimate natural shell growth. From being very low in the smallest mussels, growth was linear between about 2–10 yr of age, corresponding to about 3–20 mm length, after which it decreased with a L=32 mm. Over the linear interval, growth in the populations from 3–6 m and 10–15m depth was 3.1 and 2.2 mm · yr^-1, respectively. Meat growth showed strong annual variations mainly due to gonad production. Starving mussels could, however, while utilizing energy reserves, survive losses of up to 78% of their meat biomass. This ability of M. edulis to respire away its own biomass and its apparent tolerance of weight loss has important implication. It will drastically reduce the energy flow to destruents from mussels dying naturally, which is of special significance in the Baltic, where predators and scavengers are scarce. It enables the mussels to endure bad food conditions and buffer strong seasonal variations in food abundance, maintaining the strongly food-and space-limited Baltic M. edulis population at the carrying capacity of the area.     

Genetic and nutritional control of the gametogenic cycle in Mytilus edulis

Individuals of Mytilus edulis were collected from two populations in easterm Long Island Sound, USA, at 6 wk intervals from January 1982 to February 1983, and the hypothesis that nutrient availability controls the timing of gametogenic events in this mussel was tested using genetic variants at the Lap locus. In eastern Long Island Sound, Lap genotypes differ in the net rate of nitrogen accumulation. A highly significant difference in reproductive cycle among genotypes occurred in a population of M. edulis where Lap genotypes differ in nitrogen budget. Genotypes with lower rates of nutrient accumulation delayed the reproductive cycle by approximately six weeks relative to genotypes with high rates of nutrient accumulation. No significant difference in reproductive cycle occurred at a locality where Lap genotypes do not differ in nitrogen budget. These data indicate that nutritional status can exert significant control over the timing of gametogenesis in M. edulis. Differences in reproductive cycle among Lap genotypes provide a mechanism of non-random mating among genotypes which may result in heterozygote deficiencies at this locus.     

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.     

Clinal variation and heterozygote deficit at the lap-locus in Mytilus edulis

Electrophoretic investigations of Mytilus edulis populations on the New York and Connecticut coasts of Long Island Sound (USA) revealed a cline in the frequency of the Lap ⁹⁴-allele at a 20 km zone at the entrance to the estuary. Significant deficits of heterozygotes were found in the region of the cline and are discussed in terms of a model of mixing between oceanic mussel populations of high gene frequency and estuarine populations of low frequency. The pattern of estuarine circulation is assumed to provide a mechanical basis for isolation between oceanic and estuarine populations and to determine the position of the gene frequency cline.     

Effect of suspended bottom material on growth and energetics in Mytilus edulis

The influence of suspended, natural silt (0 to 20 mg l^-1) in addition to unicellular algal cells (Phaeodactylum tricornutum) (o to 20.000 cells ml^-1) on clearance, growth and energetics in Mytilus edulis has been studied. Clearance increased by 32 to 43% by the addition of 5 mg silt l^-1 as compared to clearance in a pure algal suspension. Ingestion and growth rate increased with algal concentration, and growth rate was further increased by 30 to 70% by the addition of 5 mg silt l^-1. A growth rate comparable to maximum natural growth rates was reached only at the highest algal concentration in the presence of 5 mg siltl^-1. Assimilation efficiency of P. tricornutum decreased from 77% at 5,000 cells ml^-1 to 52% at 20,000 cells ml^-1. In the experiments with silt added, some 20 to 30% of the assimilated organic matter originated from the suspended bottom material. Net growth efficiency increased with growth rate at a decelerating rate, approaching a maximum of about 70%. It is concluded that suspended bottom material, which is always present in M. edulis' natural habitats, serves as an additional food source, and that M. edulis depends on suspended bottom material to exploit fully its clearance potential, and to reach the maximum growth rates observed in nature.     

Resource allocation in Mytilus edulis on the shore and in suspended culture

The annual cycle of carbon and nitrogen content of the flesh of wild and cultivated mussels (Mytilus edulis L.) in Killary Harbour, Ireland, was measured over two years, starting in February, 1980 and ending in November, 1981. The carbon and nitrogen contents of mussel gametes were determined and the allometry of growth of wild and cultured mussel shells was examined with respect to length, weight and organic content. The carbon and nitrogen contents of the organic fraction of the shell were determined. These data were combined with those we had previously published on growth rate, gametogenesis and the annual cycle of ash-free dry weight (AFDW) of mussels in the same locality. Estimates were made of fecundity, reproductive effort and the partitioning of carbon and nitrogen between soma, gametes and shell. In suspended culture, cumulative production after eighteen months is equal to cumulative production after six to seven years on the shore. For comparison, partitioning of carbon and nitrogen resources between soma, shell and gonad is estimated when total cumulative production by wild and cultivated mussels is approximately equal. Differences in resource allocation are considerable. Wild mussels allocate some 57% of their carbon budget and 52% of their nitrogen budget to gamete output. In culture, mussels allocate only 22% of their carbon budget and 19% of their nitrogen budget to gamete output. It is concluded that in response to a higher production rate, cultivated mussels increase allocation of resources to somatic growth.     

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.     

Antigenic characterization of hemocyte subpopulations in the mussel Mytilus edulis by means of monoclonal antibodies

In order to characterize in more detail hemocytes from Mytilus edulis and establish links between histological and cytological featues, monoclonal antibodies (MABs) were prepared using hybridoma technology. Five MABs were identified and their reactivity patterns were analyzed with both immunoperoxidase and ultrastructural immunogold labelling. Four MAB classes were identified by their immunostaining patterns. Optical and ultrastructural examination revealed that Class I MAB reacted specifically with basophilic granulocytes (granulocytes containing small granules). Class II and III MABs differed in the intensity of their indirect immunofluorescence and degree of immunoperoxidase labelling, but all recognized both basophilic and eosinophilic granulocytes. Epitopes recognized by Class IV MABs were not always present in basophilic granulocytes, whereas they were always expressed in the eosinophilic granulocytes (granulocytes containing large granules) and could constitute differentiation antigens which progressively form during the maturation process. Molecular weights of the proteins recognized by these MABs were determined by Western blotting. It is concluded that immunostaining can identify at least three hemocyte types in M. edulis.