Ultrastructural analysis of dissolution of shell of the bivalve Mytilus edulis by the accessory boring organ of the gastropod Urosalpinx cinerea

The shell penetrating mechanism of the muricid gastropod Urosalpinx cinerea follyensis Baker was examined with reference to solubilizing effects of the secretion of the accessory boring organ (ABO) on the ultrastructural organic and mineral components of the shell of the bivalve Mytilus edulis Linné. The fine structure of shell etched by the secretion was contrasted with normal shell and shell solubilized artificially by acids, a chelating agent, and enzymes as an aid in interpreting the pattern etched by the secretion. A synoptic series of scanning electron micrographs of representative regions of the normal shell of M. edulis was prepared to serve as a standard for ultrastructural interpretation of the patterns of dissolution. Intercrystalline conchiolin of the mosaicostracal, prismatic, myostracal, and nacreous strata was dissolved as readily as the periostracum by the ABO secretion. In the prismatic region, maximum depth of dissolution of intercrystalline organic matrix occurred when long axes of prisms were approximately perpendicular to the surface being dissolved. Microscopic solubilization of organic matrix noticeably preceded dissolution of mineral crystals, revealing subsurface prisms and lamellae similar in size and form to the distinctively shaped normal shell units. After solubilization of intercrystalline conchiolin, further dissolution by the ABO secretion revealed a variety of what appeared like internal structures in prisms and lamellae. The form of these subunits varied from that of platelets to nodules in prisms and from laths to nodules in lamellae. These intracrystalline patterns of dissolution probably resulted from preferential etching of (a) soluble intracrystalline conchiolin membranes or other internal aggregates of nacrin, (b) heterogeneously distributed trace and minor elements, or (c) from both. Carriker and Williams (1978) hypothesized that a combination of HCl, chelating agents, and enzymes in a hypertonic mucoid secretion released by the ABO dissolve shell during hole boring. The similarity of patterns of dissolution etched by the ABO secretion and those produced artificially by HCl and EDTA in the present study support the hypothesis that these chemicals, or chemicals similar to them, are constituents of the ABO secretion. Lactic and succinic acids and a chitinase-like enzyme were also suggested by Carriker and Williams as possible agents in shell dissolution. Alteration of the shell surface by experimental application of these, and other, chemical agents was not sufficiently comparable to that etched by the ABO secretion to support the suggestion. Preferential dissolution of shell matrix by the ABO secretion is functionally advantageous to boring gastropods because it increases the surface area of mineral crystals exposed to solubilization and facilitates removal of shell units from the surface of the borehole by the radula.     

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     

Patterns of mtDNA diversity in North Atlantic populations of the mussel Mytilus edulis

Patterns of (female) mitochondrial DNA diversity were investigated in the blue mussel Mytilus edulis. Mytilus edulis is a ubiquitous member of contemporary North Atlantic hard-substrate communities and well represented in studies of this region. Mytilus edulis was surveyed in North America and Europe, as well as mid-Atlantic sites in Greenland, Iceland, and the Faroe Islands. Mitochondrial DNA sequences revealed considerable population structure but no monophyly of haplotypes between any major regions. Coalescent analyses suggest that migration across the Atlantic Ocean has prominently been from North American source populations and that Greenland was colonized recently and exclusively from North America. In North America, there was support for two regional groups along the North American coastline. Surprisingly, we also found evidence of recombination between some Mytilus edulis and Mytilus galloprovincialis female mtDNA sequences, particularly in northern Europe. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. An erratum to this article can be found at     

Experimental analysis of byssus thread production by Mytilus edulis and Modiolus modiolus in sediments

Laboratory experiments have been conducted on byssus thread production by two species of mussel, Mytilus edulis L. and Modiolus modiolus (L.), representing the epibyssate and endobyssate species of the Mytilidae, respectively. Mussels were placed in seven particle sizes of sediment ranging from 50 m to 16 mm for 12 d. The number of byssus threads per mussel, length of threads, number of threads per particle and size of pads were then recorded. Modiolus modiolus (endobyssate) produced more threads than Mytilus edulis (epibyssate). M. edulis produced most threads in the size range 2 to 16 mm and Modiolus modiolus in the size range 500 to 1000 m. M. modiolus produced longer threads than Mytilus edulis. Both species produced longer threads in particle sizes finer than 2 mm. M. edulis produced smaller pads than Modiolus modiolus. Both species produced larger pads in sediments coarser than 1 000 m. Results were also analysed in terms of attachment units (a thread attached to one or more particles, or a particle to which more than one thread is attached). The thread:particle ratio of the attachment units ranged from ca. 1:10 in the finest sediments to ca. 10:1 in the coarsest sediments. Mytilus edulis attached more threads to single particles (72% of attachment units) than Modiolus modiolus (37% of attachment units). M. modiolus had a wider spread of ratios in the finer particle sizes than Mytilus edulis. The significance of our results are discussed in relation to the ecology and palaeoecology of epibyssate and endobysste species.     

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.     

Potential effects of a non-indigenous predator in its expanded range: assessing green crab, <Emphasis Type="Italic">Carcinus maenas</Emphasis>, prey preference in a productive coastal area of Atlantic Canada

The non-indigenous green crab (Carcinus maenas) is an important predator on bivalve wild beds in coastal areas worldwide. This study explored size-dependent green crab prey preference on American oysters (Crassostrea virginica), blue mussels (Mytilus edulis), and soft-shell clams (Mya arenaria) in a productive coastal system of Atlantic Canada. Using two sizes of prey and three different experimental manipulations, small, medium, and large green crabs were given a choice among these three bivalves, and their daily feeding rates were monitored over the course of 3 days. For both prey sizes, green crabs showed an early feeding preference for soft-shell clams and, only as they declined in numbers, a switch toward mussels and subsequently toward oysters. We found that such changes in the timing (order) of prey preference are related to prey differences in shell thickness, a fairly reliable indicator of prey shell strength.     

Precipitation of aragonite by calcitic bivalves in Mg-enriched marine waters

To understand the relative importance of biological versus physicochemical control over biomineralization, we have tested if the chemical composition of the medium (i.e., the Mg/Ca ratio) can change the mineralogy of mollusk shells. The shells of mollusks are made of calcite and/or aragonite, which are by far the most common CaCO₃ polymorphs. Several species of bivalves with predominantly calcitic shells have been cultivated in artificial seawater with a Mg/Ca molar ratio within the range of 8.3–9.2, well above the present value for seawater (5.2). Four out of six species used (the scallop Chlamys varia, the oyster Ostrea edulis, the saddle oyster Anomia ephippium and the mussel Mytilus edulis) survived long enough to secrete significant amounts of calcium carbonate. The deposits (sometimes extensive) formed on the interior shell surfaces were predominantly aragonitic. Three individuals of C. varia also increased their length by adding new shell at the margin. Contrary to the internal shell deposits, these margins were high-Mg calcite. This implies that the marginal mantle is able to exert a more strict control on the secreted mineral phase than the mantle facing the internal shell surface. This is the first report on an in vivo experimentally forced switch in bivalve shell mineralogy, from calcite to aragonite due to a change in water chemistry.     

Genotype-dependent fecundity and temporal variation of spawning in hybrid mussel (Mytilus) populations

Mussel samples were collected at 4 to 6 wk intervals throughout 1987 from two hybridMytilus edulis/M. galloprovincialis populations, at Croyde Bay and Whitsand Bay, in southwest England. These were analyzed at two polymorphic loci which are diagnostic for allozyme differences which typifyM. edulis andM. galloprovincialis. Dry mantle weight as a function of shell length was determined for all individuals of each sample. Size-frequency data for the two populations was obtained in September 1987 and March 1988. For all genotypes at both sites, fecundity was a function of shell length, and in both populations the frequency ofM. galloprovincialis alleles was positively correlated with shell length. At both sites, allozyme genotype explained a significant amount of variation in mantle weight either when assessed as a main effect or when assessed as an interaction with shell length or time of collection. At Croyde,M. galloprovincialis mussels had greater estimated fecundity per unit length than theM. edulis mussels. Differences in the timing of spawning activity between theM. edulis and theM. galloprovincialis mussels were inferred, and these differences might act to reduce the amount of interbreeding at Croyde. At Whitsand, a reduced level of variability in the timing of spawning activity and fecundity between the genotypes was observed and explained by a higher degree of genetic mixing. Because theM. galloprovincialis mussels had (1) a greater estimated fecundity at any length, and (2) a greater mean length than theM. edulis mussels, the mean genotypic annual fecundity perM. galloprovincialis mussel was 2.8 times greater than an individualM. edulis mussel at Croyde, and 2.2 times greater than an individualM. edulis mussel at Whitsand. This evidence thatM. galloprovincialis mussels have an advantage in fecundity, and thus perhaps in fertility, taken together with the evidence thatM. galloprovincialis also has a higher viability, indicates directional selection in favour of theM. galloprovincialis phenotype. Because of the observed temporal stability of the population it seems likely that this selection is counterbalanced by a massive imigration ofM. edulis spat from neighbouring populations.     

Feeding habits and phenotypic changes in proboscis length in the southern oyster drill, Stramonita haemastoma (Gastropoda: Muricidae), on Florida sabellariid worm reefs

The southern oyster drill, Stramonita (= Thais, Kool 1987) haemastoma, is a common intertidal and subtidal predator in the southeastern United States. It uses specialized feeding structures and foraging strategies to bore holes through the shell of its bivalve prey. However, on the east coast of Florida, S. haemastoma, is common on sabellariid worm reefs constructed by the polychaete Phragmatopoma lapidosa (Walton Rocks Beach, Florida, 27°17′N, 80°12′W), a habitat where the snail’s typical prey are scarce. From 1999 to 2001, we examined the feeding habits of S. haemastoma on sabellariid reefs and the behavioral and morphological responses of S. haemastoma that accompanied switching from a diet of bivalves to sabellariids. On worm reefs S. haemastoma feeds on P. lapidosa by inserting the proboscis deep into a worm’s tube. Worm-feeding snails had longer proboscises (~3.7 times shell height) than bivalve-feeding conspecifics (~2.0 times shell height). Snails raised on different diets showed significant differences in proboscis length suggesting that the proboscis length is phenotypically plastic. Whereas typical oyster drills must bore holes for days before ingesting prey, S. haemastoma on worm reefs avoids boring and attacks and consumes worms in 15–50 min. In the laboratory, oyster drills consumed 1.7 worms per day, spending <1 h each day feeding. On sabellariid reefs, differences in feeding, handling costs, and prey value, are likely to have a significant effect on the ecology and life history of S. haemastoma in this habitat.     

Decrease of net carbon flux in two species of mussels caused by extracts of crude oil

Effects of crude oil and salinity stress on the metabolism of two common filter-feeding animals (Mytilus edulis and Modiolus demissus) have been investigated. Carbon budgets have been calculated for each species under a variety of combinations of oil content and salinity. Both reduced salinity and the presence of crude oil tend to decrease the net carbon flux for each species; stresses from each source interacted in their effects on experimental animals. Although similar responses to oil were shown by each species, Mytilus edulis appeared to be slightly more resistant to oil than Modiolus demissus.     

Variation in strength of attachment to the substrate explains differential mortality in hybrid mussel (Mytilus galloprovincialis and M. edulis) populations

Mussel samples were collected from a hybrid mussel (Mytilus galloprovincialis and M. edulis) Population at Croyde, southwest England, in January, March and May 1990. The strength of attachment of each mussel to the substrate was measured with a spring balance. A number of diagnostic characters were also recorded. These are shell lengh, width and height, mantle colour and genotype at two allozyme loci, esterase-D and octopine dehydrogenase. Multiple-regression analysis was used to assess the effect of the diagnostic characters on strength of attachment as dependent variable. Mussels possessing the relatively high shells and darker mantle colouration characteristic of M. galloprovincialis had higher values, on average, for strength of attachment than mussels resembling M. edulis. Phenotypically intermediate mussels had intermediate values for strength of attachment. The results suggest an adaptive difference which can account for reports of differential mortality acting in favour of M. galloprovincialiis.     

Shell ecophenotype in the blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) determines the spatial pattern in foraging behaviour of an oystercatcher (<Emphasis Type="Italic">Haematopus ostralegus</Emphasis>) population

When feeding on blue mussels (Mytilus edulis), oystercatchers (Haematopus ostralegus) either stab into the mollusc’s gaping valves or hammer through its dorsal or ventral shell. Whilst the selectivity of hammering and stabbing oystercatchers for specific prey morphologies has been well studied, the way in which the effects of environment on M. edulis morphology can in turn affect feeding methods of H. ostralegus is very poorly understood. Based on morphological analyses on randomly selected shells from three intertidal zones, this study failed to detect differences in morphology or distribution of dorsally and ventrally hammered shells but confirms the finding of previous authors that hammering oystercatchers select thinner mussels than stabbing birds. Additionally, we show that this difference in optimal prey morphology can lead to spatial patterns in oystercatcher feeding behaviour. Whilst at the low intertidal and higher mid intertidal zones, characterised by comparatively thick shells, most empty shells had apparently been stabbed, hammering was the dominant feeding behaviour at the lower mid intertidal zone, where shells were thinner. Preference of hammering birds for smaller mussels was not ubiquitous. Sagittal shell shape was predominantly influenced by allometric growth effects and had only minor effect on prey selection. All oystercatchers preferred less inflated mussels, with the degree of shell inflation gradually increasing with higher intertidal elevation. Our results illustrate the importance of small-scale patterns in prey ecophenotypes in determining the distribution and feeding dynamics of wading birds.     

Effect of salinity fluctuation on the osmotic pressure and Na+, Ca2+ and Mg2+ ion concentrations in the hemolymph of bivalve molluscs

Specimens of Chlamys opercularis, Modiolus modiolus, Mytilus edulis, Crassostrea gigas, Scrobicularia plana and Mya arenaria were exposed to both gradual (sinusoidal) and abrupt (square-wave) salinity fluctuations and measurements made of osmotic, Na⁺, Mg²⁺ and Ca²⁺ concentrations in the hemolymph and where applicable in the mantle fluid. In both sinusoidal and square-wave regimes fluctuating between 100 and 50% seawater (100%=ca. 32 S), the hemolymph Na⁺, Mg²⁺, Ca²⁺ and osmotic concentrations followed the concentrations of the external medium in Chlamys opercularis. The hemolymph and mantle fluid osmotic Na⁺, Mg²⁺ and Ca²⁺ concentrations of Modiolus modiolus, Mytilus edulis, Crassostrea gigas and S. plana followed those of the external medium as long as the molluscs' shell valves remained open. There were no changes in the ionic or osmotic concentrations of the hemolymph or mantle fluid of any of these species during periods of shell-valve closure. The hemolymph osmotic, Na⁺ and Mg²⁺ concentrations of wedged-open Modiolus modiolus, Mytilus edulis, C. gigas and S. plana followed those of the external medium. Hemolymph Ca²⁺ concentrations showed a damped response in C. gigas and Mytilus edulis. The hemolymph osmotic, Na⁺, Ca²⁺ and Mg²⁺ concentrations of Mya arenaria fluctuated in a similar manner to the external medium, but were damped. Wedged-open Mytilus edulis exposed to fluctuating salinity and supplied with a constant supply of 10 mM Ca²⁺ showed greater changes in hemolymph ionic and osmotic concentrations than M. edulis exposed to the same salinity fluctuation without a constant Ca²⁺ supply. Chlamys opercularis and Modiolus modiolus survived in a 50% seawater minimum sinusoidal salinity fluctuation for 10 days; wedged-open M. modiolus survived only 3 days. Burrowing had no effect on the osmotic, Na⁺, Mg²⁺ or Ca²⁺ concentrations of the hemolymph of Mya arenaria or S. plana exposed to fluctuating salinities. All of the species studied were shown to be osmoconformers.     

Filtration rate,using a new indirect technique,in thirteen species of suspension-feeding bivalves

A method for determining filtration rates in undisturbed suspension-feeding bivalves is described. Concentrations of particulate matter in the water collected in the inhalant (C _i) and exhalant (C _e) currents were estimated with an electronic particle counter. The clearance was calculated as , where Fl=flow rate through the tube collecting exhalant water. Only above critical levels of water flow (Fl) were clearances representative of filtration rates. At 10° to 13°C, the filtration rates (F, 1 h^-1) within one or two orders of magnitude of dry weight (w, g), in Cardium echinatum L., C. edule L., Mytilus edulis L., Modiolus modiolus (L.) and Arctica islandica (L.) followed the allometric equations: 4.22w ^0.62, 11.60w ^0.70, 7.45w ^0.66, 6.00w ^0.75 and 5.55w ^0.62, respectively. Five species of bivalves [Spisula subtruncata (da Costa), Hiatella striata (Fleuriau), Cultellus pellucidus (Pennant), Mya arenaria L. and Venerupis pullastra (Montagu)] filtered with the same rates as individuals of Cardium echinatum and A. islandica of equivalent soft weight. In Pecten furtivus and P. opercularis filtration rates were about twice the rates measured in individuals of Mytilus edulis of comparable body weight. The gill area in M. edulis increases with size at the same rate as the filtration rate.     

Effects of seawater pCO2 and temperature on shell growth, shell stability, condition and cellular stress of Western Baltic Sea Mytilus edulis (L.) and Arctica islandica (L.)

Acidification of the World’s oceans may directly impact reproduction, performance and shell formation of marine calcifying organisms. In addition, since shell production is costly and stress in general draws on an organism’s energy budget, shell growth and stability of bivalves should indirectly be affected by environmental stress. The aim of this study was to investigate whether a combination of warming and acidification leads to increased physiological stress (lipofuscin accumulation and mortality) and affects the performance [shell growth, shell breaking force, condition index (C_i)] of young Mytilus edulis and Arctica islandica from the Baltic Sea. We cultured the bivalves in a fully-crossed 2-factorial experimental setup (seawater (sw) pCO₂ levels “low”, “medium” and “high” for both species, temperature levels 7.5, 10, 16, 20 and 25 °C for M. edulis and 7.5, 10 and 16 °C for A. islandica) for 13 weeks in summer. Mytilus edulis and A. islandica appeared to tolerate wide ranges of sw temperature and pCO₂. Lipofuscin accumulation of M. edulis increased with temperature while the C_i decreased, but shell growth of the mussels only sharply decreased while its mortality increased between 20 and 25 °C. In A. islandica, lipofuscin accumulation increased with temperature, whereas the C_i, shell growth and shell breaking force decreased. The pCO₂ treatment had only marginal effects on the measured parameters of both bivalve species. Shell growth of both bivalve species was not impaired by under-saturation of the sea water with respect to aragonite and calcite. Furthermore, independently of water temperatures shell breaking force of both species and shell growth of A. islandica remained unaffected by the applied elevated sw pCO₂ for several months. Only at the highest temperature (25 °C), growth arrest of M. edulis was recorded at the high sw pCO₂ treatment and the C_i of M. edulis was slightly higher at the medium sw pCO₂ treatment than at the low and high sw pCO₂ treatments. The only effect of elevated sw pCO₂ on A. islandica was an increase in lipofuscin accumulation at the high sw pCO₂ treatment compared to the medium sw pCO₂ treatment. Our results show that, despite this robustness, growth of both M. edulis and A. islandica can be reduced if sw temperatures remain high for several weeks in summer. As large body size constitutes an escape from crab and sea star predation, this can make bivalves presumably more vulnerable to predation—with possible negative consequences on population growth. In M. edulis, but not in A. islandica, this effect is amplified by elevated sw pCO₂. We follow that combined effects of elevated sw pCO₂ and ocean warming might cause shifts in future Western Baltic Sea community structures and ecosystem services; however, only if predators or other interacting species do not suffer as strong from these stressors.     

Effect of tidal submersion on lysozyme activity in Mytilus edulis and Tellina tenuis

Changes in mean style weight and in lysozyme activity of the style, digestive gland, gill and mantle of Mytilus edulis and Tellina tenuis from the Clyde Sea area, Scotland, were investigated over tidal cycles in March and August, 1981. For M. edulis, significant changes occurred in the style weight, style lysozyme activity and digestive gland lysozyme activity during a 22 h period. These appear to be related to a diurnal cycle rather than a tidal cycle. Changes in the weight of the style of M. edulis may be caused by dissolution during feeding, and style lysozyme may be secreted independently of the style matrix. The activity of lysozyme in T. tenuis is unaffected by the tide, suggesting that this intertidal bivalve can feed throughout the tidal cycle.     

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.     

Induced defensive responses by the bivalve Mytilus edulis to predators with different attack modes

Predators such as crabs, whelks, and sea stars attack their bivalve prey in different ways, and predator-induced defenses are an important means of protection. The degree to which induced defenses are specific to different predators, however, remains largely unknown. In laboratory experiments (June to August 1998), we raised mussels (Mytilus edulis L.) in the presence of a drilling predator [the whelk Nucella lapillus (L.)] or a crushing predator [the crab Carcinus maenas (L.)] to determine whether induced changes in prey shell thickness, size, or shape occurred and whether changes were predator-specific. Over a 2 month period, juvenile mussels were exposed to waterborne cues from actively feeding crabs or whelks. Mussels produced thicker shell lips in response to both predators relative to control mussels raised in their absence, and the difference was significantly greater in response to whelks than to crabs. Mussels exposed to whelks showed significantly smaller increases in shell length and width and total wet weight than did mussels exposed to crabs. Thus, there may be a trade-off between shell thickness and linear shell growth and a potential delay in attaining a size refuge from predation. Received: 4 August 1999 / Accepted: 31 January 2000     

Chromium in intertidal sediments of the Clyde,UK: potential for remobilisation and bioaccumulation

Detailed surveys of intertidal sediments have been performed along the north and south shores of the Inner Clyde estuary, UK. Surface sediment data reveal significant spatial variation in Cr content and an association with major sediment characteristics and location within the estuary. Depth variation for Cr and other heavy metals cannot be explained by variation in major geochemical controls such as grain size and organic matter and highlights the impact of historical contamination on sediment quality. These elevated levels at depth may still have environmental impact through redox-reactivity, in association with iron and manganese. Sequential extraction of sediments and pore water analysis of Cr(VI) and Cr(III) provide detailed information on release potential from the sediments. The implication of Cr mobility for biota in the estuary has been assessed by the analysis of a common marine bivalve, Mytilus edulis (Blue Mussel) and a burrowing polychaete, Nereis diversicolor (rag worm) from a number of survey sites. Bioconcentration factors for Mytilus indicate that the weakly held portion of sediment Cr is available for uptake and in the case of Nereis, bioaccumulation appears to be inhibited by sediment organic matter.     

Tube-worm-sediment relationships in populations of Pectinaria gouldii (Polychaeta: Pectinariidae) from Barnegat Bay,New Jersey,USA

Pectinaria gouldii (Verrill), which lives for 1 year in Barnegat Bay, New Jersey, constructs over its lifetime a conical tube of increasingly large sand grains, regardless of surrounding sediment characteristics. However, the rate of increase of mean grain size of the tube and the population density of the worm vary with sediment type. The distribution of this species is limited by sediment composition. Worms of equal length will always have equal anterior tube apertures, although the thickness of the tube walls may be unequal. Tube surface-area, worm dry weight, and tube weight all increase as a power function of tube length. The conical shape and increasing mass of the tube impose an upper limit to worm growth, but do not interfere with worm mobility.