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     

Mitochondrial DNA heteroplasmy in European populations of the mussel Mytilus trossulus

Mitochondrial DNA (mtDNA) of Mytilus trossulus from the Gulf of Gdansk (southern Baltic) and M. edulis from Swansea Bay, South Wales, UK, collected in 1991, was studied by restriction-enzyme analysis. These two species were more similar to each other in haplotype frequencies than either was to M. galloprovincialis from Britain. M. trossulus resembles M. edulis in having a high frequency of heteroplasmy restricted to males. However, in contrast to M. edulis where restriction-site heteroplasmy predominates, in M. trossulus heteroplasmic individuals possess two genomes which differ in length by up to 3 kilobases.     

Molecular population genetics of male and female mitochondrial genomes in subarctic Mytilus trossulus

The doubly uniparental inheritance system allows for the use of two independent mitochondrial genomes for population history investigations. Under this system, two lineages of mitochondrial DNA (mtDNA) exist and males are typically heteroplasmic, having the additional, usually divergent, mitochondrial genome inherited from their male parent. This additional mtDNA typically evolves faster, potentially allowing for insight into more recent events in population history. Few studies did explore this possibility in marine mussels Mytilus showing its usefulness. Recent observations of the Mytilus trossulus mussels who have retained their native mtDNA in European waters posed the question of their origin. Are they part of a population present, but previously undetected, or is this a potentially human mediated, ongoing spread of an invasive species? To tackle this question, we amplified with species-specific primers and sequenced an approximately 1,200-bp-long fragment spanning COIII and ND2 genes from both mitochondrial genomes of mussels sampled at five locations worldwide, covering the whole M. trossulus range. The overall pattern of polymorphisms is compatible with the entirely postglacial history of the whole species, indicating a very deep bottleneck at last glacial maximum, with possible retention of the whole species in a single refugium, and the effective population size of no more than a few thousands. Both analyses of molecular variance and isolation with migration (IM) models point at the West Atlantic as the source of the European M. trossulus mussels, at least the ones who retained their native mtDNA. The hypothesis that this is an ongoing, human-mediated process was considered. To this end, comparison with the well-known case: the introduction of congeneric mussel, Mytilus galloprovincialis, from Mediterranean Sea to Asia was used. This introduction occurred within the last 100 years. The results inferred by the IM model suggest that the timing and structure of transatlantic migration of M. trossulus differs significantly from the M. galloprovincialis case: it is more than 1,000 years old and involves a much larger fraction of the ancestral population. Therefore, most likely, this invasion is not a human-mediated process.     

Ecological gradients and relative abundance of native (Mytilus trossulus) and invasive (Mytilus galloprovincialis) blue mussels in the California hybrid zone

Marine communities are experiencing unprecedented rates of species homogenization due to the increasing success of invasive species, but little is known about the mechanisms that allow a species to invade and persist in a new habitat. In central California, native (Mytilus trossulus Gould 1850) and invasive (Mytilus galloprovincialis Lamarck 1819) blue mussels and their hybrids co-exist, providing an opportunity to analyze the mechanisms that determine the distributions of these taxa. Spatial and temporal variation in temperature and salinity and the relative frequencies of these mussel taxa were examined between 2000 and 2004 at four sites in San Francisco Bay and four in Monterey Bay, which were chosen for their different positions along inferred estuarine/oceanic gradients in the hybrid zone. Mussels were genetically identified as the parent species or hybrids by amplifying regions of two species-specific loci: the adhesive byssal thread protein (Glu-5′) and the internal transcribed spacer region of ribosomal DNA (ITS 1). The proportion of M. trossulus at the eight hybrid zone sites correlated negatively with average salinity (R ²=0.60) and positively with maximal temperature (R ²≥0.72), a somewhat unexpected result given what is known about the phylogeography of this species. The proportion of M. galloprovincialis showed the opposite pattern. The proportion of hybrids was correlated neither with habitat temperature nor salinity. Genotypes of mussel populations at an additional 13 sites from Coos Bay, Oregon (latitude 43.35°N) to Long Beach, California (latitude 33.72°N), sampled at various intervals between 2000 and 2004, were also determined. This survey confirmed previous reports that the hybrid zone lies between Monterey and the Cape Mendocino region (latitudes 36.63°N–40.5°N). Within Monterey and San Francisco Bays, however, the temporal comparisons (1990s vs. 2000s) revealed abrupt changes in the proportions of the two parent species and their hybrids on annual and decadal time scales. These changes indicate that the blue mussel populations are in a highly dynamic state. The survey also showed that, regardless of habitat, M. trossulus is consistently of smaller average size than either M. galloprovincialis or hybrids.

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p53 sequence polymorphisms in late-stage leukemic Mytilus edulis are homologous with M. trossulus p53

Mytilus edulis and M. trossulus are key species in marine intertidal communities that develop a leukemia-like haemic neoplasia at increased incidences at some polluted sites. The tumor-suppressor p53, known to be involved in this disease, was proposed as a biomarker based on sequence polymorphisms in M. trossulus. In this study, p53 mRNA sequence variations in normal and diseased M. edulis were compared with those reported previously for M. trossulus for caged individuals of both species submerged in Burrard Inlet (49°20′16.57″N 123°13′51.37″) in 2007 and 2010. Thirty-seven single nucleotide polymorphisms (SNPs) were detected in M. edulis p53 that formed high linkage disequilibrium blocks associated with late-stage leukemia (p < 0.05). These were found at loci that were more homologous with M. trossulus than with healthy M. edulis p53-predicted haplotypes. Increased susceptibility to this cancer might be due to, in part, p53 alleles containing these SNPs.     

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.     

Digestion and assimilation of protein by Mytilus trossulus (Bivalvia: Mollusca) fed mixed carbohydrate/protein microcapsules

Cross-linked walled protein microcapsules have previously been developed and shown to be ingested, digested and assimilated by suspension-feeding bivalves. In this study, we examined the effect of adding carbohydrate to protein capsules on the utilization of encapsulated protein by mussels (Mytilus trossulus) in a series of in vitro enzyme incubations and an in vivo feeding experiment. Two types of carbohydrate (amylose and maltodextrin) were separately encapsulated at a carbohydrate to protein ratio of 1:1 by weight. In vitro carbohydrate and protein leakage rates from mixed carbohydrate/protein capsules were low [e.g. <4% (24h)^-1] compared to previously reported leakage rates for protein capsules [e.g. 5 to 20% (24h)^-1]. Proteolytic breakdown of mixed carbohydrate/protein capsules by mussel-style extracts was similar to that for protein capsules, when the encapsulated carbohydrate was amylose, a straight-chain (-1,4-linkage) polysaccharide; however, capsules containing maltodextrin, which is partially comprised of -1,6-linkages, were not as readily digestible in vitro. In contrast to results from in vitro enzyme assays, maltodextrin/protein capsules were utilized in vivo by mussels with similar efficiencies to those for protein and amylose/protein capsules. These findings demonstrate that utilization of microencapsulated protein by mussels cannot be accurately predicted from results of in vitro enzyme assays. Importantly, up to 50% of the dry weight of cross-linked walled protein microcapsules can be replaced with carbohydrate without significantly affecting encapsulated protein digestion and assimilation efficiencies for M. trossulus.     

Feeding responses of the bivalves Crassostrea gigas and Mytilus trossulus to chemical composition of fresh and aged kelp detritus

The chemical composition of kelps (e.g. polyphenolics) deters grazing by herbivores, but kelp detritus is potentially a source of nutrition for suspension feeders. The effects of kelp detritus derived from two species [Agarum fimbriatum Harvey and Costaria costata (Turner) Saunders] on feeding of oysters, Crassostrea gigas Thunberg, and mussels, Mytilus trossulus Gould, were examined in feeding experiments. Fresh and aged kelp particles were sequentially presented in combination with the microalga Rhodomonas lens at an initial total concentration of 5᎒^-4 ml^-1. Aging of kelp particles for 4 days in seawater significantly reduced the concentration of polyphenolics without changing the total carbon or nitrogen content. Clearance rates of both mussels and oysters were significantly lower in the presence of fresh versus aged kelp particles, and clearance rates declined overall with declining polyphenolic concentrations. Video endoscopy was used to examine feeding selectivity at the level of the gill in oysters in the same food treatments used in the clearance rate experiments. Comparison of particle composition in the water versus the pseudofeces in both oysters and mussels was also used as a measure of feeding selectivity. When presented with R. lens in combination with fresh and aged kelp particles selectivity for R. lens tended to be greater against fresh than aged particles, and there was some indication that this was stronger for A. fimbriatum than for C. costata particles. The ability to select was lower at very high polyphenolic concentrations, which may reflect poisoning of sensory binding sites. These data suggest that bivalves distinguish among particles of varying chemical composition and respond by changing their clearance rates and their selectivity.     

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     

Hybridization between the blue mussels Mytilus galloprovincialis and M. trossulus along the Pacific coast of North America: evidence for limited introgression

Two species of blue mussel, Mytilus galloprovincialis and M. trossulus, co-occur and hybridize along the Pacific coast of North America. Using a set of polymerase chain-reaction (PCR)-based genetic markers which diagnostically identify these species, we show that they are sympatric from the Cape Mendocino region to the Monterey Peninsula in northern and central California, USA. Mussels with hybrid genotypes were detected in all populations sampled in the region of sympatry, and the frequency of hybrid genotypes in individual hybrid populations ranged from 13 to 44%. Significant frequencies of first-generation backcross genotypes were detected in two individual hybrid zone populations (Berkeley and Monterey Marina) and in the hybrid zone as a whole, indicating that the potential exists for introgression between M. galloprovincialis and M. trossulus. Despite this potential, we found no evidence of advanced introgression beyond first-generation backcrosses, suggesting that gene flow between M. galloprovincialis and M. trossulus has been quite limited. The frequency of mussels with M. trossulus and hybrid genotypes declined abruptly south of Monterey Peninsula, while the frequency of mussels with M. galloprovincialis and hybrid genotypes declined precipitously north of Cape Mendocino. These abrupt genetic discontinuities indicate that this blue mussel hybrid zone is presently positioned between two prominent coastal features and there is little, if any, export of alleles from the hybrid zone into bordering parental populations. Received: 20 August 1997 / Accepted: 26 October 1998     

Distribution of male and female mtDNA lineages in populations of blue mussels,Mytilus trossulus and M. galloprovincialis,along the Pacific coast of North America

Geographic variation in mitochondrial large subunit (16S) ribosomal RNA haplotypes was examined for blue mussels, Mytilus trossulus Gould, 1850 and M. galloprovincialis Lamarck, 1819, sampled from ten sites along the Pacific coast of the USA in January of 1993. Using polymerase chain reaction/restriction fragment length polymorphism (PCR/RFLP) assays we determined haplotype frequencies for both the male and female mussel mitochondrial DNA (mtDNA) lineages. Populations from Morro Bay south to San Diego, California, contained only M. galloprovincialis male and female haplotypes, while those from Arcata Bay, California, north to Port Orford, Oregon, were fixed for M. trossulus haplotypes. Populations from Monterey Bay to Bodega Bay, California, contained a mixture of M. trossulus and M. galloprovincialis haplotypes. Overall only 2 of 97 heteroplasmic individuals had a mixed M. trossulus/M. galloprovincialis mitochondrial genotype indicating that hybridization is uncommon in the populations sampled. Further, there was no evidence of extensive introgression between these mussel taxa at the level of mtDNA. This is in contrast to previously published results which suggested the significant introgression of M. trossulus haplotypes into southern populations containing primarily M. galloprovincialis nuclear genotypes. We feel the discrepancy lies in the ability of our assays to detect haplotypes corresponding to both the male and female mtDNA lineages. Potential explanations for the lack of mtDNA introgression include, low levels of backcrossing between hybrids and parental taxa, epistatic interaction between nuclear and mitochondrial genes and the breakdown of a sex-specific inheritance pattern for mtDNA in hybrids.     

Urban Geochemistry: A Multimedia and Multielement Survey of a Small Town in Northern Europe

The concentrations and distributions of chemical elements (Ag, Al, As, Au, Ba, Bi, Ca, Cd, Co, Cr, Cu, Fe, Ga, Hg, K, La, Mg, Mn, Mo, Na, Ni, P, Pb, S, Sb, Sc, Se, Sr, Te, Th, Ti, Tl, U, V, W and Zn) were studied in till, humus and urban soil in Jakobstad, a small town (20,000 inhabitants) in W. Finland. The analyses were performed with ICP-MS after aqua regia leaching of till (n = 37), urban topsoil (n = 32) and subsoil (n = 32), and HNO3 leaching of humus (n = 37). The till and humus samples, collected at the same sites, were divided into urban and rural samples. The urban till was not significantly enriched in metals. In contrast, a majority of the elements occurred in higher concentrations in the urban than the rural humus samples. Statistical and spatial interpretations of the humus data revealed that traffic (Pb, V and Ni), metal industry (Pb, Zn, Bi, Sb and Cr), an abandoned shooting range (Pb and Sb) and other sources contribute to higher metal levels in the urban humus. The urban soil samples were collected at parks, yards, abandoned industrial sites, roadsides, etc. The topsoil samples were enriched in most elements, also by elements not enriched in the urban humus (e.g. Cd). At several sites, the concentrations far exceeded the limit concentrations for contaminated soils in Finland. A large variety of sources were identified or indicated.     

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.     

Vanadium transfer in the mussel Mytilus galloprovincialis

Radiotracers were used to study processes controlling the accumulation and elimination of vanadium in the Mediterranean mussel Mytilus galloprovincialis. Vanadium uptake rates varied inversely with both salinity and vanadium concentration in water, but were independent of temperature. After a 3 wk exposure to ⁴⁸V, the highest concentration factors were found in the byssus (1900) with much lower values computed for shell ( 70) and soft tissues (5). More than 90% of the total ⁴⁸V accumulated was fixed to shell, suggesting that uptake is primarily a result of surface sorption processes. Much of the vanadium in shell was firmly bound to the periostracum and was not easily removed by acid leaching. Food-chain experiments indicated that the assimilation coefficient for ingested vanadium is low (7%) and that the assimilated fraction is rapidly excreted from the mussel. These findings coupled with knowledge of in situ and experimentally-derived vanadium concentration-factors have allowed a preliminary assessment of the relative importance of the food and water pathways in the contamination of mussels under conditions of acute and chronic exposure. Contaminated mussels transferred to clean sea water lost ⁴⁸V at rates that depended upon temperature but were largely unaffected by either salinity or by vanadium levels in mussel tissues. Total vanadium depuration was slow and was governed by loss from a slowly-exchanging compartment with a characteristic half-time of about 100 d. Individual mussel tissues were analyzed for stable vanadium and the possibility of using these tissues, particularly the byssus, as bioindicators of ambient vanadium levels in the marine environment is also 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.     

Frostschutz-Glykoproteine bei Mytilus edulis?

Fractions of glycoproteins preventing formation of ice crystals in water were isolated from the mussel Mytilus edulis, using ion-exchange procedure and gel filtration. The protein fractions depress the freezing point of water more than would be expected from their concentrations, taking into account their molecular weights (>10 000 Daltons). It is suggested that the occurrence of such antifreeze glycoproteins contributes essentially to the mechanism of freezing resistance in the mussel.

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Mit finanzieller Unterstützung durch die Deutsche Forschungsgemeinschaft (Th 158/7).     

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.     

The musselsMytilus galloprovincialis andM. trossulus on the Pacific coast of North America

Most recent authors have called the bay mussels of the Pacific coast of North AmericaMytilus edulis Linnaeus, 1758. Thirteen samples ofedulis-like mussels were collected from California, Oregon, and Alaska, USA, in 1985, 1986 and 1987. Electrophoretic evidence from wight loci indicates that these samples consist of two genetically distinct groups, neither of which is similar toM. edulis from the Atlantic Ocean. Mussels in southern California are very similar toM. galloprovincialis Lamarck, 1819 from the Mediterranean Sea; it is probable thatM. galloprovincialis was introduced accidentally to southern California. Mussels in Oregon and Alaska are similar to those from the Baltic Sea and parts of eastern Canada; the nameM. trossulus Gould, 1850 has priority for this taxon. In central and nothern California,M. galloprovincialis, M. trossulus and their hybrids co-occur. Despite the presence of hybrids betweenM. galloprovincialis andM. trossulus, the genetic integrity which they maintain across large areas of the world warrants their recognition as two distinct species.     

Reproductive isolation between populations from Northern and Central Europe of the leaf beetle Chrysomela lapponica L.

Summary. Allopatric populations of the leaf beetle Chrysomela lapponica are known to feed upon either willow (Salicaceae) or birch (Betulaceae). This study aimed to elucidate the differentiation process of these allopatric populations. We investigated whether these allopatric populations specialized on different host plants are still able to produce fertile offspring when interbreeding. Individuals from a population in Finland (willow specialists) and one in the Czech Republic (birch specialists) were crossed in laboratory. Hybrid formation succeeded only between females from the Czech, birch specialized population and males from the Finnish, willow specialized population, while no eggs were produced by females of the willow specialists having mated males of the birch specialists. Behavioral, morphological, physiological, and chemical features of the F₁ hybrids were studied. The chemical composition of larval defensive secretion and feeding preferences of the resulting F₁ hybrids mainly showed similarities with the paternal phenotype, while the area of black coloring on the offspring’s elytra was intermediate between those of the parental elytra. F₁ hybrids did not accept the host plant (birch) of their mothers for feeding and only survived on willow. Thus, since mothers only lay eggs on birch, we found evidence for a postzygotic isolation mechanism between the individuals of the two investigated populations: when having been mated with willow specialized Finnish males, the birch specialized Czech mothers place the hybrid eggs on a plant species (birch), on which the hatching larvae cannot survive.