A hydromechanical principle for particle retention in Mytilus edulis and other ciliary suspension feeders

It is generally thought that the laterofrontal cirri of the bivalve gill act as filters that retain suspended particles in the through current and transfer the particles onto the frontal surface of the gill filaments. In Mytilus edulis calculations indicated that if water passed between the branching cilia of the cirri that are assumed to constitute the filter the pressure drop needed would amount to about 10 times the actual pressure drop across the whole gill. Thus, instead of acting as filters the laterofrontal cirri seem to move water. Presumably, the cirri together with the frontal cilia produce the water currents along the frontal surface of the gill filaments. Particle retention in the bivalve gill implies the transfer of suspended particles from the current of water about to enter an interfilamentar space into a neighbouring frontal surface current. The complex three-dimensional pattern of flow that arises where the 2 systems of current meet is characterized by steep velocity gradients. Particles that enter such steep, steady velocity gradients become exposed to transverse forces that cause the particles to migrate perpendicularly to the direction of flow. Whether particles enter the surface current, i.e. are retained, or they stay within the through current andescape, depends primarily upon particle size, and upon the steepness and height of the gradients within the boundary zone between the surface current and through current. Further studies are needed to evaluate the capacities and relative importance of this hydromechanical particle-trapping mechanism in suspension feeding bivalves. It is suggested that in downstream particle-retaining systems, e.g. on the tentacles of polychaetes and entoprocts, velocity gradients between through currents and surface currents also act as the particle-collecting mechanism.     

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     

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.     

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.     

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.     

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.     

Oogenesis in the marine mussel Mytilus edulis: an ultrastructural study

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

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).     

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.     

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.     

Food quality and season affect gene expression of the mucosal lectin MeML and particle sorting in the blue mussel Mytilus edulis

The blue mussel, Mytilus edulis, is able to use biochemical cues to sort food particles. Recently, a mucosal lectin (MeML) identified in pallial organs of M. edulis was suggested to play a role in the capture and sorting of food particles. The current study presents data obtained in 2010 showing that sorting efficiencies and MeML gene expression in M. edulis increased when mussels were either fed poor quality food or starved. In addition, results suggest that particle capture and degree of selection increased during the spawning period, although the quality of food delivered to mussels before the experiments seemed to have a stronger impact on particle-sorting efficiency. Overall, the impact of both endogenous (physiological status) and exogenous (food quality) factors were shown to affect the expression of MeML in mussel feeding organs, and the food sorting abilities.     

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.     

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.     

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.     

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.     

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.     

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     

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.     

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.