A possible hybrid zone in the Mytilus edulis complex in Japan revealed by PCR markers

The adhesive protein allele of mussels collected at 13 points in Japan from Hokkaido to Kyushu was analyzed by the polymerase chain reaction using a set of primers which amplifies a part of the nonrepetitive region of the adhesive protein gene. While most mussels exhibited a 126 bp fragment, characteristic of the pure Mytilus galloprovincialis, 55 of 64 mussels sampled at Hiura and 1 of 14 mussels at Hakodate Port exhibited 168 and 126 bp fragments. Sequence analysis of the two fragments indicated that the 168 and 126 bp fragments are almost identical to previously reported sequences in M. trossulus and M. galloprovincialis, respectively. Since the frequency of heterozygous individuals in Hiura is very high, it is unlikely that they are simple hybrids. However, it is evident that mixing of genes occurred between the two species off Hokkaido. Received: 6 September 1996 / Accepted: 9 October 1996     

The relationship between allele frequency and tidal height in a mussel hybrid zone: a test of the differential settlement hypothesis

The blue mussels Mytilus edulis L. and M. galloprovincialis Lmk. hybridize in western Europe. Within hybrid populations nuclear alleles specific to M. galloprovincialis increase in frequency with age and size. This relationship changes with tidal height; alleles from M. galloprovincialis occur more frequently high in the intertidal zone, while M. edulis alleles predominate in the low intertidal zone. We tested the hypotheses that larvae with M. galloprovincialis alleles tend to settle higher in the intertidal zone, or that mussels redistribute themselves with respect to tidal height after initial larval settlement. We sampled recently metamorphosed mussels every 2 weeks in a hybrid mussel population at Whitsand Bay in southwest England throughout the summer of 1996. We observed four cohorts of newly settled mussels. There was no evidence of differential settlement of mussels with different genotypes in connection with tidal height, or into shaded versus unshaded microsites. Therefore, we rejected the preferential settlement hypothesis. There was substantial movement of juvenile mussels in the first 4 weeks following initial settlement, but this “secondary settlement” did not result in genetic differentiation with respect to tidal height. Further, significant differences in allele frequencies were found between primary and secondary spat. This allele frequency change was in the opposite direction of that seen in the adult population, suggesting newly settled larvae may be experiencing different selective pressures than adults. We propose that the genetic structure of hybrid mussel populations with respect to tidal height is the consequence of differences in selection intensity. Received: 30 April 1999 / Accepted: 5 May 2000     

Effect of Cd on accumulation and loss of 210Po in the mussel Mytilus edulis

As a result of ²¹⁰Po's previously identified association with sulphur-rich proteins, metallothioneins could have a significant effect on the behaviour and fate of ²¹⁰Po in molluscs. Starved control and cadmium-exposed mussels, Mytilus edulis, were fed ²¹⁰Po-labelled algae (Isochrysis galbana) for 5 d and then allowed to depurate in clean sea water. Cadmium-exposed M. edulis accumulated less ²¹⁰Po in the digestive gland and the remainder of the tissue than control mussels, although this was due to a decrease in tissue weight. More than 40% of ²¹⁰Po was identified as being associated with high molecular weight and heat-treated cytosol proteins in M. edulis. Mussels in a starved state are known to recycle as much as 90% of their amino acids. It is proposed that ²¹⁰Po associated with these and other proteins is recycled, explaining why no significant loss of ²¹⁰Po was observed from the remainder of the tissue in either control or Cd-exposed mussels. Cadmium-induced metallothioneins had no effect on the distribution of ²¹⁰Po in M. edulis; <5% associated with the cytosolic fraction was considered to principally contain metallothioneins. It is suggested that ²¹⁰Po's apparent relationship with metallothioneins is coincidental rather than connected with its role in the regulation of metals. Received: 30 March 1998 / Accepted: 3 August 1998     

Oil-induced disruption of foraging behaviour of the asteroid keystone predator, Coscinasterias muricata (Echinodermata)

A prey-localization test allowed the observation of the foraging behaviour of Coscinasterias muricata in the presence of Mytilus edulis and the disruption of this behaviour following oil exposure. Asteroids were exposed to dilutions of water-accommodated fractions (WAF) of Bass Strait stabilised crude oil (control, 2%, 10%). Effects of exposure (4 d) and depuration (20 d) were quantified using circular statistical analyses. Observations suggested that disruption of behaviour might be concentration-dependent. Control asteroids successfully located mussels during tests. A small proportion (3 of 16) of asteroids exposed to 2% WAF also located the mussels, while asteroids exposed to 10% WAF did not. Following depuration, exposed asteroids recovered their chemoreception capacities. It is concluded that (1) Coscinasterias muricata is able to locate prey mussels through chemoreception, (2) exposure to oil disrupts its foraging behaviour, and (3) the effect is reversible. Received: 4 March 1998 / Accepted: 25 November 1998     

Assessment of natural selection in a hybrid population of mussels: evaluation of exogenous vs endogenous selection models

We examined natural selection within a population of marine mussels, sampled in southwestern England in June 1991, containing a high frequency of hybrids between Mytilus edulis L. and M. galloprovincialis Lmk. This system is particularly tractable for the assessment of natural selection because hybridization is common and individual mussels can be aged, allowing changes in the frequency of hybrid genotypes among age classes to be determined. We show that strong viability selection occurs among hybrid genotypes which results in the virtual elimination of M. edulis–like genotypes from the population over a period of 3 years. Recombinant hybrid genotypes are intermediate in fitness, with M. edulis–like genotypes having a lower survival rate and M. galloprovincialis–like genotypes having a higher survival rate than genotypes of mixed ancestry. Since intermediate fitness for hybrid genotypes is inconsistent with endogenous selection models we conclude that the structure and position of this hybrid zone is probably generated by exogenous selection. This pattern of selection is a recurring feature of this hybrid population and likely occurs elsewhere in the hybrid zone. Selection against M. edulis–like genotypes appears to be offset by extensive immigration of larvae dispersed from pure populations of M. edulis. Received: 14 July 1997 / Accepted: 24 February 1998     

Reproductive isolation and reproductive output in two sympatric mussel species (Mytilus edulis, M. trossulus) and their hybrids from Newfoundland

The mussels Mytilus edulis L. and M. trossulus Gould are found sympatrically in most areas of Newfoundland, with a low frequency of hybrids. To assess the potential for reproductive isolation, we sampled mussels from three sites in an eastern Newfoundland Bay from May–October 1996 to determine if there were differences in the reproductive cycles of the two species and their natural hybrids. In mussels with sheil lengths of 38–42 mm, males and females with mature gametes were dominant in June for M. edulis and hybrids, while M. trossulus showed a lower frequency of individuals with mature gametes. M. trossulus and hybrids spawned over a prolonged period (from late spring to early autumn) compared with most M. edulis individuals that spawned over a period of 2–3 weeks in July. This asynchrony in spawning activity between the two species may partially explain the low frequency of hybrids found in previous studies of these mussel populations. Female and male hybrids between M. edulis and M. trossulus showed normal gonad development, ripening and spawning, providing an opportunity for the introgression of genes between the two species. M. trossulus had a higher reproductive output than M. edulis of similar shell length, while hybrids showed intermediate values of reproductive output. M. trossulus females produced smaller eggs than either M. edulis or hybrids. Differences in reproductive traits may partially explain the maintenance of the mussel hybrid zone in Newfoundland. Published online: 13 August 2002     

Some aspects of nitrogen metabolism in Mytilus edulis: effects of aerial exposure

The effects of starvation and emersion on the ammonia effluxes of Mytilus edulis L. were studied. Both fed and starved groups showed similar patterns of efflux during re-immersion, indicating no compensation for starvation during anaerobic catabolism; this is taken as evidence that carbohydrate or lipid is the predominant energy source at the outset of anaerobiosis. Instead of an overshoot of ammonia excreted during the 4 h following re-immersion, all groups showed significantly reduced efflux rates independent of emersion duration. Such results suggest some conversion of ammonia during the recovery period, as no significant drop in haemolymph ammonia occurred at this time. The level of ammonia accumulation in haemolymph and mantle cavity fluid decreased exponentially with emersion duration, implying the stabilisation, at a low rate, of ammonia production after a relatively long period of emersion (16 to 24 h). Oxygen levels of the mantle cavity fluid dropped rapidly during emersion but never attained anoxia over 24 h of emersion, which may indicate some oxygen uptake by emersed mussels. It has been concluded that the behaviour and physiological responses shown by M. edulis to emersion appear to reflect a need for respiratory gas exchange more than a response to desiccation – presumably in the interest of energy-conservation. The reductions in ammonia efflux rate during emersion and, to a certain extent, immediately after re-immersion have also been considered here to be energy-saving strategies that illustrate the importance of ammonia in restoring normal levels of some amino acids following re-immersion. Received: 26 August 1997 / Accepted: 27 May 1999     

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     

Characterization of barnacle (Balanus eburneus and B. cenatus ) adhesive proteins

Cements from two species of barnacles, Balanuseburneus Gould and Balanus crenatus Bruguiére, were analyzed to identify the number and nature of proteins present. B. crenatus cement was composed mainly of a small peptide cross-linked into aggregates ranging in size from ˜ 3 kD to 40–50 kD. These aggregates could be reduced with 2-mercaptoethanol (2-ME) in the presence of sodium dodecyl sulfate (SDS) but only after incubation at 40 °C for 12 to 24 h and only when the cement was recently formed. Reductive alkylation of cysteine residues with 4-vinylpyridine produced a water-soluble peptide of less than 5 kD. By comparison, the cement of B. eburneus could be dissolved only partially in SDS and 2-ME when heated at 100 °C for 10 min. Five major proteins were identified by SDS-PAGE: 7, 22, 36 and 58 kD bands for which N-terminal sequence and amino acid compositions are presented; and a 52 kD band for which sequence data are given. A minor protein band of ˜ 80 kD has the same N-terminus as the 36 kD band. CNBr digests of individual proteins produced peptides for which sequence and composition data are also presented. The study was conducted during 1993 to 1995. In general, the proteins identified from B. crenatus cement are similar to those characterized from B. eburneus, and they are different in composition and sequence than those previously reported from Mytilus edulis. Received: 7 August 1996 / Accepted: 8 October 1996     

Food-particle size and selection by bivalve larvae in a temperate embayment

Epifluorescence microscopy was used to analyze the stomach contents of bivalve larvae collected in the Baie des Chaleurs (western Gulf of St. Lawrence, Canada) in order to document food-particle sizes, compare feeding among taxa, and compare the diet with the in situ phytoplankton community. Stomach contents were mainly composed of small autotrophic flagellates (<5 μm) and cyanobacteria (<2 μm), reflecting the microbial food web which characterizes these waters. More than half (55%) of all veligers examined contained algal cells of 5 to 15 μm, whereas only 3% had cells of 15 to 25 μm. Differences in the size ranges of ingested algal cells among similar-sized larvae of different species suggests that veligers actively selected food particles. Among the smallest veligers (185 to 260 μm), scallops (Placopecten magellicanus) and mussels (Mytilus edulis) ingested more <5 μm and 5 to 15 μm algae than clams (Mya arenaria). Among larger veligers (261 to 405 μm), clams contained significantly more <5 μm cells than mussels, whereas mussels contained significantly more 5 to 15 μm algae than clams. Algal cells of 15 to 25 μm were preferentially ingested by mussel veligers. Feeding also differed between different-sized veligers within taxa, i.e. the smallest clam veligers ingested fewer of 5 to 15 μm algae than the larger size classes. Mussel veligers ingested significantly more 15 to 25 μm and fewer <5 μm cells as their size increased. The dominance of ultraplankton in the nearshore waters of Baie des Chaleurs and in the stomach contents suggests that veliger larvae may be an important export path for carbon produced by small phytoplankton. Received: 17 July 1996 / Accepted: 20 September 1996     

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.     

Selective feeding in shellfish: size-dependent rejection of large particles within pseudofaeces from Mytilus edulis, Ruditapes philippinarum and Tapes decussatus

Mytilus edulis L., Ruditapes philippinarum (Adams & Reeve) and Tapes decussatus L. were fed particles of the same shape (spherical), the same density (2.1 g cm⁻³) and the same chemical composition (SiO₂), but which varied in diameter from 5 to 37 μm. Findings obtained at different particle concentrations (mean ± SD) of 51 ± 2, 105 ± 18 and 171 ± 17 mg l⁻¹ invariably indicate that significant proportions of all particles with diameters larger than from between 7.5 and 22.5 μm were preferentially rejected as pseudofaeces. We define the preferential ingestion index (PII) as the ratio between average particle volume in pseudofaeces and average particle volume in food. Whatever the particle concentration or the species, this PII was always statistically higher than 1. Irrespective of particle concentration, PII values in M. edulis were lower than in T. decussatus (averages of 1.2 and 2, respectively). PII values in M. edulis were also lower than in R. philippinarum maintained at particle concentrations above 171 ± 17 mg l⁻¹. We suggest that preferential size-dependent rejection of larger particles could be of significant adaptive value in the natural environment, either if there are large inorganic particles, or if the average organic content of smaller particles is higher. Received: 11 January 1997 / Accepted: 8 March 1997     

A molecular approach to the ecology of a mussel (Mytilus edulis – M. trossulus) hybrid zone on the east coast of Newfoundland, Canada

Mytilus trossulus Gould and M. edulis L. coexist in mixed populations in Atlantic Canadian waters. Although there is evidence that the two species hybridize in natural populations and that hybrids produce progeny through backcrosses, no study of the microgeographic distribution of the two forms and their hybrids has been made. Here we examine subtidal samples of mussels taken in July 1997 from two locations in eastern Newfoundland (Canada) and from wave-exposed and protected environments within each location. Shell lengths ranged from 15 to about 90 mm. Mussels were classified as pure forms or hybrids (F₁, F₂ and from backcrosses) based on four diagnostic markers, two allozyme loci (Mpi and Est-D) and two nuclear PCR-based DNA markers (ITS and Glu-5). In addition, a PCR-based mtDNA marker (COIII) was used to characterize the distribution of mtDNA mitotypes among pure and hybrid individuals. There were differences in the proportions of pure M. edulis and M. trossulus between sites and between environments at one location. M. trossulus was the predominant species at one of the two exposed sites. In all four samples, M. trossulus was also the predominant form among small individuals. The frequency of hybrids was 26% overall and did not differ among samples. Hybrids consisted mostly of backcrosses that were M. trossulus-biased among small mussels and M. edulis-biased among large ones. We conclude that both intrinsic genetic factors and extrinsic environmental factors influence the relative frequency of M. edulis, M. trossulus and their hybrids. Received: 29 June 1998 / Accepted: 4 November 1998     

Assimilation of 210Po by the mussel Mytilus edulis from the alga Isochrysis galbana

Marine invertebrates are thought to accumulate ²¹⁰Po primarily from their food. In this study, a pulse-chase methodology was used to examine the assimilation and depuration of ²¹⁰Po by Mytilus edulis from the common marine alga Isochrysis galbana. The digestion of ²¹⁰Po from I. galbana occurred via a biphasic process, characteristic of a rapid (extracellular) and slow (intracellular) digestion typical of marine bivalves. The mantle/gill and foot have no known digestive role, yet their ²¹⁰Po specific activities increased after 24 h. It is proposed that this increase in ²¹⁰Po specific activity was related to ²¹⁰Po being incorporated into these tissues from ²¹⁰Po assimilated from I. galbana during extracellular digestion. It is proposed that the linear loss of ²¹⁰Po previously accumulated by control mussels was related to the continual state of renewal and replacement of cellular proteins, with ²¹⁰Po turnover and metabolism governed by protein turnover and metabolism. M. edulis' assimilation efficiency of ²¹⁰Po from the ²¹⁰Po-labelled alga was calculated to be 17.2 ± 2.1%, and thus similar to that of Ag, Cd, Co, Se and Zn by bivalves from other marine algae species. It is proposed that the assimilation efficiency of ²¹⁰Po is a function of protein assimilation. Received: 27 August 1998 / Accepted: 3 September 1999     

Influence of cadmium accumulation and dietary status on fatty acid composition in two colour forms of shore crabs, Carcinus maenas

 The effects of cadmium exposure and dietary status on cadmium accumulation, fatty acid (FA) content and profiles were investigated in two colour forms of the shore crab Carcinus maenas. Groups of shore crabs were either starved or fed with blue mussels, Mytilus edulis, during a 40 d exposure period to 2 or 6 μM Cd²⁺ (as CdCl₂). Starved green individuals accumulated more cadmium in haemolymph and hepatopancreas than did red crabs and green crabs fed during the experiments. In the red colour form, no difference in cadmium accumulation was observed between starved and fed individuals. In both colour forms, hepatopancreas contained more FA than gills and muscle. The FAs often present in the largest amounts in the tissues were 16:0, 16:1ω7, 18:1ω7, 18:1ω9, 20:4ω6, 20:5ω3 and 22:6ω3. However, saturated (SAFA) and mono-unsaturated fatty acids (MUFAs) were dominant in hepatopancreas, whereas poly-unsaturated fatty acids (PUFAs) were dominant in gills and muscles. At the beginning of the experiment, the total FA content in the hepatopancreas was 111.6 mg g⁻¹ (dry weight) for red crabs and 78.4 mg g⁻¹ for green shore crabs. During the experiment, however, the FA content decreased in red crabs. This decrease was more pronounced for starved individuals than for fed individuals. Also, the decrease in FA content was more pronounced in crabs exposed to 6 μM cadmium compared to crabs exposed to 2 μM or crabs not exposed to cadmium. No change in FA content was observed in green shore crabs, irrespective of diet and cadmium exposure. For both colour forms, no change in FA content was observed for gills and muscle. In red crabs, a decrease was observed for all FAs in the hepatopancreas. This decrease, however, was more pronounced for SAFAs and MUFAs than for PUFAs, indicating that the metabolism of FAs during starvation and cadmium exposure is selective. The experiments indicate that green colour forms of shore crabs are more tolerant of natural stress such as starvation and anthropogenic stress, e.g. cadmium exposure, than are red colour forms of shore crabs. Received: 23 September 1999 / Accepted: 29 April 2000     

Temporal variation of lysosomal capacities in relation to susceptibility of mussels, Mytilus edulis, to summer mortality

We examined the temporal variation of lysosomal enzyme activities and lysosomal membrane stability in two stocks of blue mussels (Mytilus edulis L., 1758) from the Magdalen Islands which differ in their susceptibility to summer mass-mortality. The activity of lysosomal enzymes reflects the autolytic capacity which could be used during the transfer of reserves between storage and reproductive tissues or for mobilisation of reserves for energy requirements. The peak of lysosomal enzyme activities in mantle tissue in June 1992 probably was related with autolysis of storage cells to support maturation of the developing gametes. A second peak of lysosomal enzyme activity which occurred in mid- August of 1992 and 1993 may help to cover energetic requirements of maintenance metabolism during this period of high temperatures and reduced food quality. Measures of the destabilization of lysosomal membranes in the digestive gland confirmed that mussels showed a significant stress response in mid-August. Mussels from the stock with a higher susceptibility to summer mortality underwent a longer period of stress and accumulated less glycogen between spawning and the stressful period in mid-August than mussels from the more resistant stock. In conclusion, our field observations demonstrate high activities of lysosomes not only in spawning periods but also during stressful periods of high temperatures and reduced food quality. Received: 23 September 1997 / Accepted: 11 June 1998     

Genetic structure of hybrid mussel populations in the west of Ireland: two hypotheses revisited

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels, the blue mussel, Mytilus edulis, and the Mediterranean mussel, M. galloprovincialis. Results from an Irish study in the 1980s, using partially diagnostic allozyme markers, indicated that mussels higher up the shore were more galloprovincialis-like than those lower down. In this study we set out to test two hypotheses: (a) recruits arriving on the shore are composed of genetically distinct cohorts that settle preferentially at different levels on the shore, and maintain genetic distinctiveness into adulthood; (b) recruits are genetically homogeneous, but once settled they diverge genetically over time, due to within-habitat site specific-selection. The diagnostic Me 15/16 DNA marker was used to analyse the genetic composition of newly-settled spat recruiting to artificial substrates, which were placed at two-week intervals from May–October 2002, on the mid- and low shore areas of two exposed sites in Galway Bay. Adult mussels were also collected on each sampling date. Results did not support the preferential settlement hypothesis, i.e., the genetic composition of primary settlers (≤ 500 μm) was similar between tidal heights and shores. Neither was there evidence of post settlement selective mortality, as adults were genetically similar to settling spat. In spat and adults the frequency of the M. galloprovincialis allele was high (0.56–0.80), due to high frequencies of M. galloprovincialis (> 37%) and hybrid (> 33%) genotypes, and correspondingly low frequencies of the M. edulis genotype (< 11%). Adult mussels from a nearby sheltered estuarine site, while significantly different to exposed shore mussels, still had low frequencies of the M. edulis genotype (< 17%), indicating no apparent advantage for the genotype in this environment. There are indications that the genetic composition of mussels may be changing on the Atlantic coasts of Ireland.     

Size-dependent, spatial and temporal genetic variation at a leucine aminopeptidase (LAP) locus among blue mussel (Mytilus galloprovincialis) populations along a salinity gradient

Biochemical genetic variation at a leucine aminopeptidase (LAP) locus is related to salinity variation in several marine bivalve molluscs. This paper details an investigation of the Long Island Sound model of LAP selection (LAP genotype-dependent mortality occurring among newly settled Mytilus edulis mussels) carried out in 1997 among three M. galloprovincialis mussel populations along the salinity gradient of Wellington Harbour, New Zealand. Significant LAP genotypic heterogeneity was observed at the LAP locus between small (<25 mm shell length) and large (>25 mm shell length) M. galloprovincialis from Petone and Eastbourne (the two sites experiencing the greatest salinity variation), whereas genotypic heterogeneity was not significantly different between small and large mussels from Seatoun (the site experiencing the least salinity variation). The Lap ³ allele decreased in frequency and the Lap ⁴ allele increased in frequency at Petone and Eastbourne, whereas the Lap ³ and Lap ⁴ allele frequencies remained effectively constant at Seatoun. Both these findings are consistent with the Long Island Sound model of selection. At all three locations, the Lap ^3,3 genotype decreased in frequency from small to large mussels, whereas the Lap ^3,4 genotype increased in frequency from small to large mussels. All other LAP genotypes occurred at low frequencies (<0.10) at all three locations and showed no evidence of frequency change from small to large-size mussels nor evidence of clinal change among the three locations. These genotype frequency data possibly indicate that the Lap ^3,3 genotype is at a selective disadvantage compared to the Lap ^3,4 genotype at all three locations, and that this selective disadvantage is related to the extent of salinity variation which exists at each location. Further investigation is required before it can be determined if the Long Island Sound model of selection best describes the size-dependent and location-specific changes in LAP allele and genotype frequencies along this salinity gradient. Comparison of the population genetic structure at the LAP locus in 1995 and in 1997 revealed a profound change from heterozygote excesses to heterozygote deficiencies for all three M. galloprovincialis populations. The reason for the change is unknown, but the change indicates that population genetic structure at the LAP locus is highly variable in time, but consistent in space, among these M.␣galloprovincialis populations. Received: 5 February 1998 / Accepted: 27 May 1998     

Uptake and depuration of cesium in the green mussel Perna viridis

 The accumulation and depuration of Cs in the green mussels (Perna viridis) commonly found in the subtropical and tropical waters were studied under the laboratory conditions using radiotracer techniques. Following an initial rapid sorption onto the mussel's tissues, uptake of Cs exhibited linear patterns over a short exposure time (8 h) at different ambient Cs concentrations. The concentration factor was independent of ambient Cs concentration. The calculated uptake rate and initial sorption constant of Cs were directly proportional to the ambient Cs concentration. The calculated uptake rate constant from the dissolved phase in the mussels was as low as 0.026 l g⁻¹ d⁻¹. Uptake rates of Cs in the mussels were inversely related to the ambient salinity. Uptake increased about twofold when the salinity was reduced from 33 to 15 ppt. The effect of salinity on Cs uptake was primarily due to the change in ambient K⁺ concentration. The uptake rate decreased in a power function with increasing tissue dry weight of the mussels, although the initial sorption was not related to the mussel's body size. The efflux rate constant of Cs in the mussels was 0.15 to 0.18 d⁻¹, and was the highest recorded to date among different metals in marine bivalves. The efflux rate constant also decreased in a power function with increasing tissue dry weight of mussels. A simple kinetic model predicted that the bioconcentration factor of Cs in the green mussels was 145, which was higher than measurements taken in their temperate counterparts. The bioconcentration factor also decreased in a power function with increasing tissue dry weight of mussels. Received: 27 October 1999 / Accepted: 16 June 2000     

Effects of oil pollution on the freezing tolerance and solute concentration of the blue mussel Mytilus edulis

Exposure of blue mussels (Mytilus edulis L.) to oil-polluted sea water for 7 d in the laboratory did not affect their body fluid concentrations of inorganic ions and free amino acids. Mussels exposed to-4°C for 12 h did not freeze, whereas freezing occurred invariably in mussels exposed to-7°C or lower temperatures for the same period. Following freezing at-10°C, oil polluted mussels resumed normal activity considerably more slowly than unpolluted ones, but oil-polluted as well as unpolluted mussels showed normal activity 3 h after thawing. Freezing at-15°C was invariably lethal to individuals of both groups. One possible explanation of the delayed recovery of oil-polluted mussels frozen at-10°C may be that oil components had become concentrated to toxic levels as the amount of solvent water diminished during freezing.