Burrow morphology,tube formation,and microarchitecture of shell dissolution by the spionid polychaete Polydora websteri

Burrow morphology, detrital tube formation within the burrow, and ultrastructure of the surface of the burrow of Polydora websteri Hartman, 1943, are described in the shells of Crassostrea virginica (Gmelin) and of Mytilus edulis Linné. Observations were made on live worms living normally in artificial preparations of polished shell covered with transparent plastic film. P. websteri is capable of settling wherever there are crevices in shell surfaces, and slowly penetrates the shell forming a U- or flask-shaped cavity. A detrital tube is formed within the burrow from detritus collected and pressed in place by the worm. During this process, the modified setae of the 5th setiger are thrust against the wall of the inner tube and help to keep the diameter of the inner tube constant throught its length. Individuals of P. websteri burrow into the shell when placed on preparations of polished shell of Mytilus edulis under plastic film. As they do so, they secrete a viscous fluid which dissolves interprismatic and interlamellar organic matrices and then dissolves the exposed crystals. Etched prisms and lamellae reveal a complex pattern of internal dissolution. Enlargement of the burrow by the worm thus takes place by chemical dissolution of shell and also probably by flushing of loosened partially dissolved prisms and lamellae from the tube. Some shell fragments become incorporated in the walls of the detrital tube. There is no evidence of setal abrasion on the shell walls of the burrow even at the ultrastructural levelUniversity of Delaware College of Marine Studies Contribution No. 99.     

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.     

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     

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.     

Chemical elements in the aragonitic and calcitic microstructural groups of shell of the oysterCrassostrea virginica: A proton probe study

We report the results of a study with proton-induced X-ray emissions (PIXE) of the distribution and concentration of 15 chemical elements (Na to Sr in periodic chart) in four microstructural and two mineralogical regions of shell of rapidly growing adult oysters [Crassostrea virginica (Gmelin)]. Hatchery-raised oysters were grown in Broadkill Estuary, Delaware, USA, for 16 wk in summer 1978. Their valve edges were filed as a marker, and the oysters were replaced in the estuary, where they grew rapidly. Shell deposited after marking had a normal microstructure and mineralogy after a narrow zone of disturbance. After 17 d oysters were sacrificed, and different mineralogical and microstructural regions of valves of three oysters, and parts of valves of two oysters, were analyzed for the elements Na to Sr. Quadrats (2.5 × 0.45 mm) included three calcitic groups (prismatic, foliated, chalky) and one aragonitic (myostracal) microstructural group; four quadrats were on the exterior and five on the interior of right and left valves. Inhalant and exhalant margins of valves and ground right and left valves of one oyster were also analyzed. Elemental chemistry of different regions of shell varied among the three microstructural groups within the single calcitic polymorph, between aragonitic and calcitic regions, and between exhalant and inhalant margins of the valves. Elements were most concentrated in the prismatic region of the right valve. Element concentrations were similar in ground right and left valves, except for higher levels of Si, Fe, and Zn in the right valve (corresponding to their high contents in prismatic shell) and of Cl in the left valve (reflecting high concentration in chalky shell, abundant in this valve). Na, Mg, Cl, Cr, Cu, Zn and Br were more concentrated in prismatic than in foliated shell. Chalky shell contained higher concentrations of Na than did prismatic shell, and high concentrations (but lower than in prismatic shell) of Mg, Cl, Ti, Mn, Fe, Zn and Br. Element concentration in myostracum was approximately the same as, or lower than, in foliated shell, except for Sr, which was higher than that in any other shell group. In the right valve most elements were concentrated in inhalant margins, and on the left valve, in exhalant margins. With increased weathering of the exterior surface of prismatic shell, Mg, Si, and Mn increased in concentration and Na, Al, Cl, Ti, Cr, Fe, Br, and Sr decreased.     

Biodegradation of shells of the black pearl oyster,Pinctada margaritifera var. cumingii,by microborers and sponges of French Polynesia

The composition, distribution and infestation sequence of organisms that destroy the commercially valuable shells of the black oyster Pinctada margaritifera var. cumingii Jameson, 1901 were studied. Three ecologically different groups of boring (euendolithic) organisms were identified: (1) phototrophic boring microorganisms (cyanobacteria, Hyella caespitosa, Hyella sp., Mastigocoleus testarum, Plectonema terebrans, and green algae, Phaeophila dendroides, Ostreobium quekettii); (2) heterotrophic boring microorganisms (fungi, Ostracoblabe implexa); (3) filter-feeding boring organisms (sponges, Cliona margaritiferae, C. vastifica). The phototrophic endoliths dominate the external pristmatic region of the shell, whereas the valuable interior nacreous region is attacked mainly by heterotrophs. Boring patterns reflect in part the shape and behaviour of the organisms and in part the structural properties of the shell, and inflict different types of damage. Infestation starts with microbial borers, which prepare the conditions for later invasion by more damaging clionid sponges. The infestation begins always at the apex, the oldest part of the shells, from which the periostracum is often removed by natural attrition or by cleaning procedure. The rate of bioerosion in 1 yr-old hatchery shells is 36 times higher than in natural populations.     

X-ray diffraction study of the first larval shell of Ostrea edulis

X-ray powder diffraction was used to study the calcification of the first larval shell of Ostrea edulis (sampled in Limski kanal, Istria, Adriatic Sea in April 1986) from the trochophore stage to the veliger larvae (prodissoconch I), and development of the latter up to several days postfertilization (prodissoconch II). In the first stage, only the amorphous component is present (periostracum and organic matrix). The beginning of shell formation is manifested by the appearance of calcite (up to 1–4% of the total vol.) and then aragonite (2 to 7%). In a later stage of the veliger larvae the fraction of calcite decreases, as well as the fraction of the amorphous component, while the fraction of aragonite rapidly increases. In the prodissoconch II stage, aragonite is dominant, with a very small amount of amorphous component and traces of calcite. In contrast, the valves of the adult O. edulis are composed mainly of calcite, with traces of aragonite.     

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.     

X-ray diffraction study of calcification processes in embryos and larvae of the brooding oyster Ostrea edulis

X-ray powder diffraction was used to study shell calcifications of the oyster Ostrea edulis, sampled in the Limski Kanal, Istria (Adriatic Sea), in May 1992. All the developmental stages were followed, from the embryonic stage through the transition between the trochophore and veliger larva (prodissoconch I and II) and later, after swarming, the pelagic free-swimming larval stages, up to their settlement and attachment (from the D-shaped to the fully formed pediveliger larva), and finally during metamorphosis and juvenile stages (dissoconch). In the first gastrula stage, only an amorphous tissue is present (a periostracum and organic matrix). The beginning of shell formation (at the end of gastrulation) in early trochophores is manifested by the appearance of calcite (up to 1–7% of total volume) and then aragonite (about 1%). In the later stage of the veliger larva the fraction of calcite decreases as well as the amorphous fraction, while the fraction of aragonite rapidly increases. In the prodissoconch II stage and during the whole pelagic period aragonite is dominant, accompanied by a very small amorphous fraction and traces of calcite. The shell mineral composition does not change until metamorphosis, whereupon the fraction of calcite rapidly increases and the fraction of aragonite decreases. The postmetamorphic valves of the juvenile and adult oyster consist mainly of calcite, except the resilium and myostracum which remain aragonitic, possibly as a continuation of the inner layer of the larval shell. Received: 28 May 1997 / Accepted: 1 July 1997     

Shell structure and behaviour related to cementation in oysters

Shell microstructure and mantle behaviour relating to shell cementation was studied on adult oysters,Crassostrea gigas, Saccostrea mordax, andS. kegaki (collected from Shirahama, Wakayama Prefecture, Japan, in 1989 and 1990). At the place of cementation, the prismatic structure of the outermost shell layer is modified to a significant structure. This structure, named the ridge-and-furrow structure, consists of calcified ridges a few microns wide separated by furrows, both arranged parallel to the shell growth direction. The furrows are ultimately filled by shell material. The prismatic and ridge-and-furrow structures gradually merge in a transitional area where an intermediate type of the two structures occurs. The small size of the crystal units of the ridges and furrows is due to the close distribution of crystal seeds, especially close to pre-existing ridges. This is the basis of the difference between the ridge-and-furrow and the prismatic structures and also makes the former structure functional for cementation, in contrast to the latter. At the site of ongoing shell cementation, the mantle margin adpresses the shell margin onto the substrate. Experiments show that this pressing action is essential for cementation and probably also for the formation of the ridge-and-furrow structure. Even the right valve, which oysters nerve use to cement in natural conditions, forms the ridge-and-furrow structure and cements to the substrate if the pressing action of its mantle margin is induced under artificial conditions. Behavioural changes probably led oysters to switch from byssal attachment to cementation within a short time span when they acquired their cementing habit.     

An exploratory study with the proton microprobe of the ontogenetic distribution of 16 elements in the shell of living oysters (Crassostrea virginica)

Study of the chemical composition of shell of exoskeletonous organisms in the past has required the sacrifice of the organism. Because the beam of the proton microprobe is relatively nondestructive and analyzes the surface layer of the shell, organisms do not have to be killed. The present paper presents results of a preliminary experiment in which distribution of elements (Na to Sr) in shell of living juvenile oysters, Crassostrea virginica (Gmelin), was studied in situ with a proton microprobe at monthly intervals for four months. The relative concentration of 16 elements was measured in the newly deposited prismatic edge of the right valve of three oysters reared in controlled laboratory conditions. Na, Mg, Al, Si, S, Cl, K, Ca, Ti, Cr, Mn, Fe, Cu, Zn, Br, and Sr were detected in concentrations as low as a few parts per million relative to the concentration of standards added to pure CaCO₃. Concentration of elements varied nominally among shells of the three individual oysters and in their successive ontogenetic stages. Fluctuations in concentration of Na, Mg, S, Cl, Ca, Mn, Fe, Cu, and Zn were generally similar in the two normally growing oysters, but differed from those in the oyster that stopped growing. Trends in concentration of Al, Si, and Sr were similar in the three oysters: those of Br were variable. Relative concentrations of Na, Cl, S, Mn, Fe, and Zn increased slightly with age of oysters, that of the other elements stayed relatively constant. Concentration of most elements was higher in shell than in seawater. Variable concentrations, especially of Na, Cl, and Si in valve edges, tend to support the hypothesis of earlier workers that separate mineral phases are present as impurities entrapped within the shell during calcification.     

In situ mapping of growth lines in the calcitic prismatic layers of mollusc shells using X-ray absorption near-edge structure (XANES) spectroscopy at the sulphur K-edge

The microstructure and composition, including chemical speciation of sulphur (S), of two mollusc shells were investigated using a combination of scanning electron microscopy, X-ray absorption near-edge structure spectroscopy (XANES) and electron probe microanalysis (EPMA). The shell of Pinna is composed of monocrystalline, and Pinctada, of polycrystalline, calcite prisms separated by organic-rich walls. Sulphur speciation information from XANES spectra using a scanning X-ray microscope showed that the protein S content of the interprismatic walls is higher than the SO₄ content, whereas the reverse is true for the intraprismatic structures. High-spatial-resolution XANES maps for the different S species across adjacent calcite prisms confirm their distinctive distributions in the molluscan shells and illustrate the presence of narrow, submicron transverse growth features. On a larger scale, a series of wider growth zones, incorporating the submicron zones, are aligned parallel to each other and cross-cut many calcite prisms. EPMA element maps for magnesium (Mg) and S demonstrate that these growth increments are compositionally zoned, comprising alternating layers of high mineral (Mg-rich) and high organic (S-rich) components. Additionally, these maps confirm that the organic interprismatic walls have lower Mg and higher S than the intraprismatic structures.     

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     

Measurement of calcium carbonate deposition in molluscs by controlled etching of radioactively labeled shells

Rates of calcium carbonate removal from shell pieces of Argopecten irradians (Say) and Mercenaria mercenaria (L.) were dependent on the type of etching fluid used and not shell origin. Etching was uniform over the entire shell surface, but surface morphology differed with etching fluids. Peak radioactivity was in early eluant fractions of shells etched immediately after radioactive labeling, and in later fractions when individuals were placed in isotope-free seawater after labeling. The etching technique can measure growth during the labeling period and subsequently by estimating the amount of calcium in fractions prior to the radioactive peak. Geometry of shell layers influenced the pattern of radioactivity seen in fractions. Peak location varied inside and outside the pallial line of individual M. mercenaria. A significant portion of the inorganic carbon used in shell formation by M. mercenaria was derived from metabolic CO₂.     

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.     

Phylogeographic study of the dwarf oyster, Ostreola stentina, from Morocco, Portugal and Tunisia: evidence of a geographic disjunction with the closely related taxa, Ostrea aupouria and Ostreola equestris

Despite the economic importance of oysters due to the high aquaculture production of several species, the current knowledge of oyster phylogeny and systematics is still fragmentary. In Europe, Ostrea edulis, the European flat oyster, and Ostreola stentina, the Provence oyster or dwarf oyster, are both present along the European and African, Atlantic and Mediterranean, coasts. In order to document the relationship not only between O. stentina and O. edulis, but also with the other Ostrea and Ostreola species, we performed a sequence analysis of the 16S mitochondrial fragment (16S rDNA: the large subunit rRNA-coding gene) and the COI fragment (COI: cytochrome oxidase subunit I). Oysters were sampled from populations in Portugal (two populations), Tunisia (two populations) and Morocco (one population), identified as O. stentina on the basis of shell morphological characters. Our data supported a high degree of differentiation between O. stentina and O. edulis and a close relationship between O. stentina and both Ostrea aupouria (from New Zealand) and Ostreola equestris (from Mexico Gulf/Atlantic). The status of this geographic disjunction between these closely related species is discussed. Furthermore, although identified in a separate genus Ostreola by Harry (Veliger 28:121–158, 1985), our molecular data on O. stentina, together with those available for the other two putative congeneric species, O. equestris and Ostreola conchaphila, would favour incorporation of Ostreola in Ostrea. Finally, a PCR-RFLP approach allowed the rapid identification of O. edulis and O. stentina.     

Genetic variation among life-history stages of mussels in a<Emphasis Type="Italic"> Mytilus edulis–M</Emphasis>.<Emphasis Type="Italic"> trossulus</Emphasis> hybrid zone

The mussel species Mytilus edulis L. and M. trossulus Gould coexist and hybridize throughout a large area that includes the north coast of Maine and Atlantic Canada. Previous studies provided genetic evidence for limited hybridization between the two species for mussels >15 mm. The present study used two genetic markers (ITS, Glu-5) to examine the genetic composition of early life-history stages by sampling veliger and pediveliger larvae, juveniles (<2.0 to 15.0 mm shell length) and adults (>15 mm shell length) in Trinity Bay, Newfoundland, during three consecutive years (1995–1997) to determine if differential mortality limits the relative abundance of hybrids. The relative frequency of the two species and the different hybrid genotypes was similar among the larvae, juveniles and small adult mussels. The double hybrid genotype (F₁-like) was the rarest genotype observed. There was no evidence for differential mortality during the early life-history stages, and factors limiting production of hybrids appear to operate before the late larval stage. The observed frequency of hybrids is probably due to a combination of pre- and postzygotic reproductive isolating mechanisms operating early in the mussels life history. M. trossulus dominated the early life-history stages, possibly due to a higher population density and a greater reproductive output than M. edulis. Differential mortality may explain the observed decrease in frequency of M. trossulus and increase in frequency of M. edulis with increasing shell length. A similar frequency of hybrid mussels from larvae to the size class of 55 mm shell length may indicate a rate of mortality intermediate between the two parental species. The M. edulis–M. trossulus hybrid zone appears to be maintained by reproductive isolating mechanisms limiting the production of hybrids and life-history differences that allow the two species to coexist.Communicated by J.P. Grassle, New Brunswick     

Heterozygosity and growth in transplanted mussels

Growth comparisons were made involving mussels (Mytilus spp.) collected from five different localities in Britain in 1980–1981. Two of the localities, Mumbles, South Wales, and Bude, Southwest England, have pure populations of M. edulis and M. galloprovincialis, respectively. The other three, Whitsand Bay, Southwest England, Croyde Bay, Southwest England and Robin Hood's Bay, Northeast England, have hybrid populations with both M. edulis and M. galloprovincialis ancestry. To make growth comparisons, mussels from different populations were mixed in oyster nets and transplanted to three different localities. After periods of transplantation varying between several months and one year, growth was assessed by measuring increase in shell length or dry body weight. Starch-gel electrophoresis was used to assay variation in the transplanted mussels at three allozyme loci partially diagnostic for M. edulis and M. galloprovincialis. The results provide evidence of small growth differences between populations and between allozyme genotypes within populations. These differences accounted for no more than a few percent of the total variation in growth between mussels. Statistically significant results were obtained, but were frequently found not to be reproducible. There is no clear evidence of a growth difference between M. edulis and M. galloprovincialis. Allozyme heterozygotes appear to have growth rates intermediate between allozyme homozygotes; this study thus fails to provide evidence for overdominance with respect to growth rate.     

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     

An electron-microscope study of periostracum formation in some marine bivalves. II. The cells lining the periostracal groove

The histology and ultrastructure of the epithelial cells of the outer and middle mantle folds of the bivalves Mytilus edulis (L), Cardium edule (L), Macoma balthica (L) and Nucula sulcata Bronn are described. The cells lining the inner face of the outer fold exhibit a prominent granular endoplasmic reticulum, and Golgi complexes which are concerned with the elaboration of granules and vesicles eventually incorporated into the periostracum. A gradual reduction in the protein synthetic apparatus occurs towards the tip of the fold. Within the cells, it is proposed that the ovoid inclusion bodies are lysosomes and that they control the rate of secretion. The cells of the middle fold are cuboidal in appearance. Those of M. edulis and N. sulcata exhibit prominent granules, whereas those of C. edule and M. balthica possess vesicles. The cells of M. edulis differ from the others in possessing stout bundles of filaments, which occupy large areas of the cell and constitute a cell web. The cells of the epithelium in all cases do not appear to be implicated in periostracum formation.