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.     

Physiological energetics of cultivated mussel (Mytilus edulis) populations in two Scottish west coast sea lochs

Physiological responses of suspended cultured mussels, Mytilus edulis L., in two Scottish sea lochs (Lochs Etive and Leven) were investigated in their native and transplanted environments, after 15 d, 4.5 mo and 1 yr acclimatization, during the main growing season of May to September 1992. These measurements were integrated by means of the balanced energy equation, and scope for growth was calculated to assess the performance of each stock. Transplanted mussels showed clear signs of stress during the first 15 d after transfer, with low clearance rates and energy retention and high rates of respiration and nitrogen excretion. There were significant differences in some of the physiological responses (clearance, respiration and excretion rates), scope for growth and growth efficiency between the native populations, with the responses of mussels in Loch Etive being more favourable than those in Loch Leven. With newly transplanted mussels after 15 d acclimatization, almost all these measurements also differed significantly from native mussels in their host site and the original stocks but, with the exception of ammonia excretion rates, all the variables of cross-transplanted mussels after 4.5 mo acclimatization were the same as those of the native stock at the host site. Good agreement between observed long-term growth rates and estimated scope for growth suggests that, like growth rate, differences in physiological responses are mainly controlled by environmental factors; i.e., stock or origin had no significant influence on variations in physiological response (except ammonia excretion). The results further indicate that scope for growth estimated during the main growing season can be used to assess the actual growth rate and to compare sites for on-growing.     

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     

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.     

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.     

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.     

Minimal predation upon meiofauna by endobenthic macrofauna in the Exe Estuary,south west England

Field manipulation experiments were performed in the Exe Estuary, south west England, in October 1988, to investigate the importance of the meiofaunamacroinfauna trophic link in benthic trophodynamics. Four hypothetically meiofauna-predacious endobenthic macrofauna species were selected for manipulation using the criteria of high abundance and different modes of feeding: Cerastoderma edule (filter-feeder), Nereis diversicolor (omnivorous scavenger), Ophelia bicornis (sand-ingester), Scrobicularia plana (deposit-feeder). Enclosures constructed from plastic tubing, 63 m nylon monofilament mesh and galvanized steel were deployed, containing adult members of these taxa at densities raised to approximately four times that of the surrounding sediment. The experiments ran for 12 tidal cycles. Differences in phyletic meiofaunal abundance between treatment and control enclosures at the termination of the experiment were assessed using both uni- and multivariate dataanalysis techniques. Only two univariate significant differences (p<0.05) existed for the N. diversicolor treatment and two for the S. plana treatment. Annelida, Turbellaria and copepod nauplii were the only meiofauna taxa affected. No univariate significant differences were recorded for either the C. edule or O. bicornis treatments. Multidimensional scaling ordination of the data revealed no consistent changes in community composition between treatments and controls. It is concluded that the experiments provide evidence of minimal predation by macroinfauna upon meiofauna.     

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.     

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).     

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.     

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.     

Thermal tolerance of Mytilus edulis of Eastern Australia

The tolerance and resistance parameters of Mytilus edulis were investigated using probit analysis based on the dosage-mortality response curve. The effect of thermal acclimation on incipient lethal temperatures for a range of 5° to 25°C are described. The ultimate upper incipient lethal temperature is 28. 2°C. Changes in the thermal resistance parameters with size, photoperiod and salinity are shown to be statistically significant. The temperature of incapacitation of the mussel is defined in relation to the critical thermal maxima. A correlation was found between latitude and both temperature of incapacitation and temperature of spawning in the laboratory.     

Seasonal cycles in the carotenoid content in Mytilus edulis

The carotenoid content of Mytilus edulis was studied over a period of 3 years from 1965 to 1967; a seasonal cycle was observed which appeared to be controlled by the spring bloom of phytoplankton, and the sexual cycle of the mussel. Infestation by the copepod Mytilicola intestinalis and hours of exposure in the tidal cycle were found to contribute to variations in carotenoid content. Maturation of the gonads and spawning occurred in the late spring and early summer of the period studied. In addition, a second sexual phase was recorded in October 1967. The possible effect of exposure on the time of spawning is discussed. Lack of food affects both carotenoid content and the maturation of the gonads. M. edulis kept at 10 °C with little food showed no seasonal variation in the carotenoid content when compared with animals fresh from the shore. Lack of food resulted in regression in gonadal tissue after a period of time, and an absence of mature and spawning individuals.     

The Mytilus edulis species complex in southwest England: effects of hybridization and introgression upon interlocus associations and morphometric variation

Loglinear analysis of electrophoretic data from two hybrid Mytilus edulis x galloprovincialis populations in southwest England revealed non-significant associations between genotypes at four allozyme loci, each of which is partially diagnostic for differences between the two taxa. Significant non-random genotypic associations within the context of the non-significant model involved all four assayed loci equally, consistent with their occurrence in a relatively tight linkage group. Multivariate analyses were used to examine electrophoretic variation from the two hybrid populations, and morphometric variation in the hybrid populations and in four allopatric (two M. edulis and two M. galloprovincialis) populations from western Europe While the number of hybrid mussels is high at both sites (22% at Croyde, 53% at Whitsand) the two taxa have largely maintained the genetic differences which exist between them in allopatry. However, morphological differences between the taxa have been eroded for mussels within the hybrid zone, whereas these differences are quite pronounced for mussels from allopatric populations. It is proposed that each taxon within the genus maintains its genetic identity, despite high dispersal potential, widespread hybridization, and high levels of introgression, as a result of adaptation to different environments. The worlwide occurrence of all four Mytilus hybrid zones at ecotones between recognized biogeographical provinces which are characterized by differences in temperature and salinity is consistent with such an interpretation.     

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.     

No. 2 fuel oil compounds in Mytilus edulis

Mytilus edulis contaminated by a brief 2-d exposure to a slick from a No. 2 fuel oil spill in the Cape Cod Canal, Massachusetts, USA were sampled six times during an 86-d post-spill period to study the rate of release of fuel oil compounds under field conditions. Detailed measurements of compounds by high resolution glass capillary gas chromatography and quantitative glass capillary gas chromatography-mass spectrometry-computer systems analyses provided a more comprehensive examination of release rates of different types of compounds. Biological half-lives were calculated for selected compounds for the first 21 d during which the release rates were exponential. Typical half-lives were n-alkanes, 0.2–0.8 d; pristane, 1.5 d; C-2 (dimethyl or ethyl) napthalenes, 0.9 d; methyl phenanthrenes, 1.7 d. Changes in relative ratios of C-2 phenanthrenes during the release period were observed. The evidence available to date strongly supports the role of molecular weight and accompanying properties of water solubility as the main controlling factors in the rate of release of fuel oil compounds by M. edulis. However, the data for the rapid release of n-alkanes and C-2 phenanthrenes also indicate molecular type and molecular configuration as additional key factors. The data from this study are compared and contrasted to data from short term experimental studies in the laboratory, longer term studies from chronic exposure conditions, and data from two other oil spills with longer term exposure.     

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.     

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.     

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     

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.