Mussels flexing their muscles: a new method for quantifying bivalve behaviour

We employed a novel technique to quantify how blue mussels Mytilus edulis react to predation risk in their environment by quantifying mussel gape using a Hall sensor attached to one shell valve reacting to a magnet attached to the other. Change in gape angle per second (CHIGA) versus gape angle plots resulted in a distribution with a boundary, which defined the maximum CHIGA of a mussel at all gape angles. CHIGA boundary plots for all individual mussels were similar in form. However, the CHIGA boundary increased in extent with mussel length (maximum CHIGA for mussel valve closures for mussels 2.98 and 79.6 mm long were −1.5 and −11°s⁻¹, respectively), showing that larger mussels opened and closed most rapidly. Mussel extract added to the seawater, a factor believed to signal predation, caused mussels to close significantly faster than otherwise (P < 0.001). This approach for assessing how mussels react to their environment indicates that mussel response to predation is graded and complex and may well indicate animal-based assessments of the trade-off between effective feeding and the likelihood of predation.     

Beat frequency of cilia in the branchial basket of the ascidian Ciona intestinalis in relation to temperature and algal cell concentration

To elucidate the effects of temperature and algal cell concentration on pumping of water in the ascidian Ciona intestinalis a number of different experiments were performed. Beat frequency of the lateral cilia in the openings of the branchial sac was measured in intact specimens using a microprojection objective and a monochrome CCD video camera. At constant low algal cell concentration, beat frequencies increased linearly with temperature from 4.0 Hz (±0.5) at 7.4 °C to 13.6 Hz (±1.6) at 20.1 °C. At a constant temperature of 15 °C, beat frequency decreased with increasing algal cell concentration from approximately 3000 to >10 000 Rhodomonas sp. cells ml⁻¹. The decrease was observed both in experiments where the ascidians had been acclimated to a fixed algal cell concentration and in experiments with changing concentrations. Effect of algal cell concentration on squirting/siphon closure and flow velocity in the exhalent siphon was measured using a thermistor. At low algal cell concentrations, flow velocity in the exhalent siphon was stable, apart from a few short squirts. At very high algal cell concentrations, the flow velocity was reduced and much less stable, with prolonged squirting. The effect of gut content on filtration was studied in experiments with specimens acclimated to high algal cell concentrations. Results showed a close relation between gut clearance and filtration rate. From the experimental results and a qualitative analysis of the Ciona-pump it was concluded that the ciliary beat frequency is proportional to the water flow through the sea squirt and that changes in pumping caused by temperature or algal cell concentration are under nervous control or governed by enzyme kinetics, rather than being a result of physico-mechanical properties, i.e. pump efficiency versus flow resistance, of the ascidian pump. Received: 6 October 1997 / Accepted: 8 October 1998     

Application of Laser Doppler Spectroscopy (LDS) in determining swimming velocities of motile phytoplankton

Pumping rates in Mytilus edulis L. were measured by means of a constant-level-tank method, in which hydrostatic pressure differences between inhalant and exhalant water levels were recorded by means of a laser beam reflected from a tethered mirror floating on the water surface. Hydrostatic pressure gradients were determined to ±0.05 mm H₂O or better. The developed technique of directly measuring pumping rates in mussels is not subject to the artefacts of other methods. The pumping rates measured in M. edulis were substantially higher than those previously determined by means of direct techniques, but similar to the maximum filtration rates, as obtained by means of two indirect techniques, i.e. about 50 ml min^-1 for a 0.15 g dry weight mussel. Positive hydrostatic pressures drastically affected water pumping. The pumping rate decreased linearly with increasing hydrostatic pressures towards a maximum pump pressure between 3 and 5 mm H₂O. Negative pressures only affected the pumping rate slightly or insignificantly, except when the mussels were exposed to rapidly increasing negative pressures. Under this condition a shunt was presumably established between the inner demibranchs, allowing water to bypass the gills.     

The invasion and subsequent die-off of <Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis> in Langebaan Lagoon,South Africa: effects on natural communities

The alien mussel Mytilus galloprovincialis invaded sand banks in Langebaan Lagoon on the west coast of South Africa in the mid-1990s. However, by 2001 these beds had completely died off, with only empty shells and anoxic sand remaining. In an effort to prevent the re-settlement of this aggressive invader, all dead mussel shells were then cleared. This study considered the impacts of the invasion and subsequent die-off on natural benthic communities. Community composition differed significantly between non-invaded and invaded areas (ANOSIM, R = 0.685 and P < 0.01) as the physical presence of mussel beds created a new habitat that promoted invasion by indigenous rocky-shore species. This dramatically increased faunal biomass from 1,132.9 g m⁻² ± 3,454.7 SD to 53,262.4 g m⁻² ± 23,052.6 SD and species richness from 38 to 49 species. Following the die-off of the mussel beds, communities remained significantly different between non-invaded areas and those in which mussel shells remained (ANOSIM, R = 0.663 and P < 0.01). Species richness was significantly greater in non-invaded areas (18 species) than in uncleared areas with remnant shells (four species) (Kruskal–Wallis ANOVA H _2,36 = 10.8964 and P = 0.032), as the previously dominant rocky-shore species became smothered by sediment and the compacted shells formed an impermeable layer excluding sandy-shore burrowing organisms. After the shells were cleared, 50% of the sandy-shore species associated with non-invaded areas returned within 5 months, but community structure still remained significantly different to non-invaded areas (ANOSIM, R = 0.235 and P > 0.05). Invasion thus dramatically altered natural communities and although the subsequent removal of the dead mussel shells appears to have aided recovery, community composition remained different from the pre-invasion state after 5 months.     

Phytoplankton selection by mussels, and diarrhetic shellfish poisoning

To better understand the dynamics of diarrhetic shellfish poisoning (DSP) contamination a field study was carried out on the feeding behavior of Mytilus galloprovincialis (Lmk) during an important DSP outbreak. The study was focused on the relationships between phytoplankton in seawater and algal cells, or their remnants, in mussel stomachs. During the period studied, M. galloprovincialis seemed to feed selectively on dinoflagellates rather than diatoms. Further selection was observed among different dinoflagellate genera, a preference for the genus Dinophysis being particularly evident. In addition, mussels seemed to open the thecae of Dinophysis cells and digest them more easily than other dinoflagellates. Due to the high variability of the results of phytoplankton analysis in the mussel stomachs, no correlation was found between the abundance of Dinophysis species in the mussels' stomachs and the content of okadaic acid plus dinophysistoxin-1 in their digestive glands, evaluated with an ELISA assay. Conversely, the presence of Dinophysis fortii (the main DSP-causative agent in the area studied) in integrated net samples of the whole water column and the toxin content of the digestive glands presented similar temporal trends. Received: 24 March 1997 / Accepted: 10 December 1997     

Inorganic mercury and methylmercury in surface sediments and mussel tissues from a microtidal lagoon (Bizerte, Tunisia)

The aim of this study was to investigate the distribution of mercury compounds in marine sediments and mussel tissues collected in the lagoon of Bizerte, Tunisia, during two seasons (summer and winter). Inorganic mercury (Hg²⁺) concentrations in sediments were found to be highly variable, ranging from 0.04 nmol.g¹ to 3.22 nmol.g⁻¹ (dry weight) with a mean value of 0.52 nmol.g⁻¹. Anthropogenic sources of Hg²⁺, most probably metallurgy or tire production industries, have been evidenced. The mean concentration of monomethylmercury (MeHg⁺) in the surface sediments is 2.32 pmol g⁻¹ ranging from below the detection limit (0.45 pmol.g⁻¹) to 14.6 pmol.g⁻¹. No significant variation was observed between winter and summer seasons for both mercury species concentration in the sediments. The Hg²⁺ concentrations in mussel tissues are also variable, ranging from 0.007 to 1.347 nmol.g⁻¹ (dry weight). The mean concentration is 0.70 nmol.g⁻¹. In these tisssues, Hg²⁺ is generally the major compound, making up ca. 88% of total mercury concentrations. However, methylmercury concentrations are significant and homogeneous, ranging from 62 to 121 pmol.g⁻¹ (mean 96 pmol.g⁻¹). The results suggest that a fraction of the inorganic mercury load in the sediments of the lagoon undergoes methylation pathways. MeHg⁺ produced is assimilated in the mussels more readily than Hg²⁺.     

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.     

Clearance rates of the great scallop (<Emphasis Type="Italic">Pecten maximus</Emphasis>) and blue mussel (<Emphasis Type="Italic">Mytilus edulis</Emphasis>) at low natural seston concentrations

Great scallop, Pecten maximus, and blue mussel, Mytilus edulis, clearance rate (CR) responses to low natural seston concentrations were investigated in the laboratory to study (1) short-term CR variations in individual bivalves exposed to a single low seston diet, and (2) seasonal variations in average CR responses of bivalve cohorts to natural environmental variations. On a short temporal scale, mean CR response of both species to 0.06 μg L⁻¹ chlorophyll a (Chl a) and 0.23 mg L⁻¹ suspended particulate matter (SPM) remained constant despite large intra-individual fluctuations in CR. In the seasonal study, cohorts of each species were exposed to four seston treatments consisting of ambient and diluted natural seston that ranged in mean concentration from 0.15 to 0.43 mg L⁻¹ SPM, 0.01 to 0.88 μg L⁻¹ Chl a, 36 to 131 μg L⁻¹ particulate organic carbon and 0.019 to 0.330 mm³ L⁻¹ particle volume. Although food abundance in all treatments was low, the nutritional quality of the seston was relatively high (e.g., mean particulate organic content ranged from 68 to 75%). Under these low seston conditions, a high percentage of P. maximus (81–98%) and M. edulis (67–97%) actively cleared particles at mean rates between 9 and 12 and between 4 and 6 L g⁻¹ h⁻¹, respectively. For both species, minimum mean CR values were obtained for animals exposed to the lowest seston concentrations. Within treatments, P. maximus showed a greater degree of seasonality in CR than M. edulis, which fed at a relatively constant rate despite seasonal changes in food and temperature. P. maximus showed a non-linear CR response to increasing Chl a levels, with rates increasing to a maximum at approximately 0.4 μg L⁻¹ Chl a and then decreasing as food quantity continued to increase. Mean CR of M. edulis also peaked at a similar concentration, but remained high and stable as the food supply continued to increase and as temperatures varied between 4.6 and 19.6°C. The results show that P. maximus and M. edulis from a low seston environment, do not stop suspension-feeding at very low seston quantities; a result that contradicts previous conclusions on the suspension-feeding behavior of bivalve mollusks and which is pertinent to interpreting the biogeographic distribution of bivalve mollusks and site suitability for aquaculture.     

The filtration rate of Mytilus edulis and its dependence on algal concentration,measured by a continuous automatic recording apparatus

A new apparatus for long-term, continuous automatic measurements of filtration rates in suspension-feeding organisms is described. As the concentration of algae in the experimental medium is diminished by the filter-feeding activity of the experimental animals, algal suspension is automatically added, thus keeping the algal concentration constant. In this way, accurate determinations of filtration rates in relation to particle concentration are made possible. For determination of filtration rates in the common mussel Mytilus edulis L., individuals of different body size (shell length 8.5 to 56.5 mm) were used. Within the range of 10x10⁶ to 40x10⁶ cells of Dunaliella marina/l, mussels of the same body size filter-out approximately the same amount of algae at high or low concentrations. A low algal concentration is counterbalanced by a corresponding higher filtration rate. Within the range of body size (W=dry weight of tissues) and algal concentrations used, the filtration rate (F) follows the general allometric equation F=a·W ^b, where a and b are constants at specific experimental conditions. At a temperature of 12 °C, the values obtained for a are 2410 at a concentration of 20x10⁶, and 1313 at a concentration of 40x10⁶ Dunaliella cells/l; correspondingly, the filtration rates of a mussel of 1 g dry-tissue weight are 2410 ml/h and 1313 ml/h. b, the slope of the regression line (0.73 to 0.74), is independent of algal concentration. However, examination of all known measurements reveals that, most probably, the general allometric equation is an oversimplification; in large individuals there is a more pronounced decrease in filtration rate. The relationship between filtration rate, body size of mussels, and algal concentrations used is discussed.This work was made possible through a research grant from the Deutsche Forschungsgemeinschaft in connection with the program Litoralforschung — Abwässer in Küstennähe.     

Deep-sea caging of the mussel Mytilus galloprovincialis: Potential application in ecotoxicological studies

We experimented with caging the Mediterranean mussel (Mytilus galloprovincialis) at various depths for 69 d to measure basic physiological parameters, histological response and bio-accumulation of contaminants in a deep-sea contaminated area. In preliminary experiments, we demonstrated, under artificial pressure conditions, the ability of mussels Mytilus galloprovincialis to tolerate rapid immersion (at a speed of up to 120 m min^?1). In situ experiments were performed using submerged lines enabling mussels to be maintained at depths ranging of 40–1550 m with survival rates ranging from 80 to 38%, respectively. No significant differences in condition indexes were observed between treated and control specimens. However, histological observations demonstrated a clear reduction in thickness of the digestive epithelium with increasing depth exposure. By determining the contaminants in caged mussels, we found the following values for chromium accumulation: 27.4 μg g^?1 dry weight at 580 m depth and 9.8 μg g^?1 dry weight at 1550 m. Selected stations were located downstream of an industrial effluent at 420 m. The biological and environmental consequences of deep-sea contamination demonstrate the suitability of caged mussels for monitoring contaminant accumulation.     

Sand and wave induced mortality in invasive (<Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis>) and indigenous (<Emphasis Type="Italic">Perna perna</Emphasis>) mussels

The ability of an invasive species to spread in a new locality depends on its interaction with the indigenous community and on variation in time and space in the environment. The Mediterranean mussel Mytilus galloprovincialis invaded the South African coast 30 years ago and it now competes and coexists with the indigenous mussel Perna perna. The two species show different tolerances to wave and sand stress, two of the main environmental factors affecting this intertidal community. P. perna is more resistant to hydrodynamic stress than M. galloprovincialis, while the invasive species is less vulnerable to sand action. Our results show that mortality rates of the two species over a period of 6 months had different timing. The indigenous species had higher mortality than M. galloprovincialis during periods of high sand accumulation in mussel beds, while the pattern reversed during winter, when wave action was high. A negative correlation between sand accumulation and attachment strength of the two mussels showed that sand not only affects mussel mortality through scouring and burial, but also weakens their attachment strength, subjecting them to a higher risk of dislodgement. Here we underline the importance of variations in time and space of environmental stress in regulating the interaction between invasive and indigenous species, and how these variations can create new competitive balances.     

Age,growth and population structure of <Emphasis Type="Italic">Modiolus barbatus</Emphasis> from the Adriatic

Age, growth and population structure of Modiolus barbatus from Mali Ston Bay, Croatia were determined using modal size (age) classes in length frequency distributions, annual pallial line scars on the inner shell surface, internal annual growth lines in shell sections of the middle nacreous layer and Calcein marked and transplanted mussels. The length frequency distributions indicated that M. barbatus attain a length of ∼40 mm in 5–6 years indicating that a large proportion of the population in Mali Ston Bay is <5 years old. Some mussels of ∼60 mm were predicted to be 14 years old using the Von Bertalanffy growth (VBG) equation. Up to the first 6 pallial line scars were visible in young (<6 years) mussels but in older shells the first scars became obscured by nacre deposition as the mussel increased in length and age. The age of the older shells (>6 years) was determined from the middle nacreous lines in shell section, which formed annually in winter between February and March; the wider dark increments forming during summer (June to September). The oldest mussel, determined from the middle nacreous lines, was >12 years, with the majority of mussels aged between 3 and 6 years of age. The ages of mussels ascertained using the growth lines were not dissimilar to the ages predicted from the length frequency distributions. Age at length curves produced using modal size class data were not different from the data obtained using the pallial scar rings and internal growth lines. Taken together these data suggest that M. barbatus attains a length of 40 and 50 mm within 5 and 8 years, respectively. Eighty one percent of individual M. barbatus injected with a Calcein seawater solution (300 mg Calcein l⁻¹), into their mantle cavity successfully deposited a fluorescent line, which was visible in suitably prepared shell sections under ultra violet light. Incorporation of Calcein into the mussel shells was seasonally variable with the lowest frequency of incorporation in mussels marked in February and recovered in May. Seasonal shell growth was observed with significantly higher growth rates in mussels marked in May and removed in August (ANCOVA, F _3,149 = 23.11, P < 0.001). Mussels (∼18 to 22 mm) marked in May and recovered in August displayed maximal growth rates of >2.5 mm month⁻¹ compared with a mean mussel growth rate of 1.2 ± 0.6 mm month⁻¹. At other times of the year mussel shell growth ranged from immeasurable to 1.48 mm month⁻¹.     

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

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

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.

---

High reproduction of <Emphasis Type="Italic">Calanus finmarchicus</Emphasis> during a diatom-dominated spring bloom

Feeding, egg production, hatching success and early naupliar development of Calanus finmarchicus were measured in three north Norwegian fjords during a spring bloom dominated by diatoms and the haptophyte Phaeocystis pouchetii. Majority of the copepod diet consisted of diatoms, mainly Thalassiosira spp. and Chaetoceros spp., with clearance rates up to 10 ml ind⁻¹ h⁻¹ for individual algae species/groups. Egg production rates were high, ranging from ca 40 up to 90 eggs f⁻¹ d⁻¹, with a hatching success of 70–85%, and fast naupliar development through the first non-feeding stages. There was no correlation between the egg or nauplii production and diatom abundance, but the hatching success was slightly negatively correlated with diatom biomass. However, the overall high reproductive rates suggested that the main food items were not harmful for C. finmarchicus reproduction in the area, although direct chemical measurements were not conducted. The high population egg production (>1,20,000 eggs m⁻² d⁻¹) indicated that a large part of the annual reproduction took place during the investigation, which stresses the importance of diatom-dominated spring phytoplankton bloom for population recruitment of C. finmarchicus in these northern ecosystems.     

Particle sorting in bivalves: in vivo determination of the pallial organs of selection

Benthic particle feeders are exposed to a food supply varying in both quantity and quality. Previous studies have shown that bivalve molluscs deal with such fluctuating particle regimes in a variety of ways, including adjustments in pumping and ingestion rates, and selective rejection of non-nutritive particles as pseudofeces. The actual site of particle selection within the pallial cavity, however, has remained a topic of speculation. During August 1995 and January and August 1996, we exposed the oysters Crassostrea virginica (Gmelin) and C. gigas (Thunberg), and the mussel Mytilus trossulus Gould to a mixture of ground, aged Spartina alterniflora Loisel and similar-sized phytoplankton at three concentrations (10³, 10⁴, 10⁵ particles ml⁻¹). We then examined the ctenidia and labial palps by means of endoscopy and sampled, in vivo, the particulate material from various ciliated tracts, and analyzed the samples with a flow cytometer. We found that in oysters, the ctenidia are responsible for particle sorting, whereas the labial palps play an accessory role in particle selection, or function to control the volume of material to be ingested. In mussels, however, the ctenidia play little role in particle selection and simply transport particulate matter to the palps for further processing. We suggest that selection by the ctenidia of oysters is a function of their architecture (plicate, heterorhabdic). Received: 4 June 1997 / Accepted: 23 January 1998     

Growth and development of nauplii and copepodites of the estuarine copepod Acartia tonsa from southern Europe (Ria de Aveiro, Portugal) under saturating food conditions

A temperature-dependent growth model is presented for nauplii and copepodites of the estuarine calanoid copepod Acartia tonsa from southern Europe (Portugal). Development was followed from egg to adult in the laboratory at four temperatures (10, 15, 18 and 22°C) and under saturating food conditions (>1,000 μg C l⁻¹). Development times versus incubation temperature were fitted to a Belehradek’s function, showing that development times decreased with increasing incubation temperature: at 10°C, A. tonsa need 40.3 days to reach adult stage, decreasing to 8.9 days when reared at 22°C. ANCOVA (homogeneity of slopes) showed that temperature (P<0.001) and growth phase (P<0.01) had a significant effect on the growth rate. Over the range of temperatures tested in this study, highest weight-specific growth rates were found during naupliar development (NI–NVI) and varied from 0.185 day⁻¹ (10°C) to 0.880 day⁻¹ (22°C) with a Q ₁₀ equal to 3.66. During copepodite growth (CI–CV), the weight-specific growth rates ranged from 0.125 day⁻¹ (10°C) to 0.488 day⁻¹ (22°C) with a Q ₁₀ equal to 3.12. The weight-specific growth rates (g) followed temperature (T) by a linear relationship and described as ln g=−2.962+0.130 T (r ²=0.99, P<0.001) for naupliar stages and ln g=−3.134+0.114T (r ²=0.97, P<0.001) for copepodite stages. By comparing in situ growth rates (juvenile growth and fecundity) for A. tonsa taken from the literature with the temperature-dependent growth model defined here we suggest that the adult females of A. tonsa are more frequently food limited than juveniles.     

Larval settlement and metamorphosis of the mussel <Emphasis Type="Italic">Mytilus galloprovincialis</Emphasis> on different macroalgae

Settlement of mussels is commonly associated with macroalgae. The effects of 19 macroalgal species on the settlement and metamorphosis of pediveliger larvae of the mussel Mytilus galloprovincialis were investigated in the laboratory. Settlement and metamorphosis inducing activities of macroalgae Chlorodesmis fastigiata and Ceramium tenerrimum collected each month during the period between January 2005 and April 2006 were also investigated. Furthermore, C. fastigiata and C. tenerrimum were subjected to various treatments to investigate the roles of bacteria and diatoms on the algal surface in the induction of larval settlement and metamorphosis of M. galloprovincialis and the characteristics of the cues in these two macroalgae. Pediveliger larvae of M. galloprovincialis settled and metamorphosed in high percentages on Cladophora sp., Chlorodesmis fastigiata, Centroceras clavulatum, and Ceramium tenerrimum, all of which were filamentous in morphology. Macroalgae that were cylindrical, phylloid, flabellate, palmate and pinnate all showed low (<8%) percentages of post-larvae but four other filamentous macroalgae also had low mussel larval settlement, suggesting that chemical factors may also be involved. Seasonal variation had no effect on inductive activities of C. fastigiata and C. tenerrimum. Treatment of C. fastigiata and C. tenerrimum with formalin, ethanol and heat resulted in the significant decrease or loss of their inductive activities. Survival of algal cells within treated macroalgae also decreased significantly. Treatment of the two macroalgae with antibiotics and GeO₂ reduced the numbers of bacteria and diatoms on their surface but did not affect their inductive activities, indicating that the cue was produced by macroalgae and not by coexisting bacteria and diatoms. However, conditioned water and crude extracts of these two macroalgae did not induce larval settlement and metamorphosis. Thus, larvae of M. galloprovincialis settled and metamorphosed on specific filamentous macroalgae. The chemical cues produced by C. fastigiata and C. tenerrimum were susceptible to ethanol and heat treatments and were not recovered in the conditioned water nor in the extracts. The finding that inactive C. tenerrimum can be produced from culturing its apical segments provides a new tool to elucidate the chemical cue(s) from macroalgae through manipulation of their culture conditions.     

Hydrodynamic stress and habitat partitioning between indigenous (Perna perna) and invasive (Mytilus galloprovincialis) mussels: constraints of an evolutionary strategy

The ability of a mussel to withstand wave-generated hydrodynamic stress depends mainly on its byssal attachment strength. This study investigated causes and consequences of different attachment strengths of the two dominant mussels species on the South African south coast, the invasive Mytilus galloprovincialis and the indigenous Perna perna, which dominate the upper and the lower areas of the lower balanoid zone, respectively and co-exist in the middle area. Attachment strength of P. perna was significantly higher than that of M. galloprovincialis. Likewise solitary mussels were more strongly attached than mussels living within mussel beds (bed mussels), and in both cases this can be explained by more and thicker byssal threads. Having a wider shell, M. galloprovincialis is also subjected to higher hydrodynamic loads than P. perna. Attachment strength of both species increased from higher to lower shore, in response to a gradient of stronger wave action. The morphological features of the invasive species and its higher mortality rates during winter storms help to explain the exclusion of M. galloprovincialis from the low shore. The results are discussed in the context of the evolutionary strategy of the alien mussel, which directs most of its energy to fast growth and high reproductive output, apparently at the cost of reduced attachment strength. This raises the prediction that its invasive impact will be more pronounced at sites subject to strong but not extreme wave action.     

Toxic metal health risk by mussel consumption

Seafood is a major dietary food worldwide. However, seafood consumption by humans can induce health risk because seafood may be contaminated by various pollutants. The mussel Mytilus galloprovincialis is widely distributed in the coastal waters of Montenegro, SouthEast Adriatic Sea. Here, Zn, Fe, Cu, Ni, Cd, Pb, As, and Hg contents in M. galloprovincialis from ten sites were analyzed to investigate health risks associated with the consumption of wild and cultivated mussels. Since there is a lack of data on the mussel consumption rate in Montenegro, the amount of mussels that can be ingested weekly over a lifetime with no risk of negative health effects was calculated using provisional tolerable weekly intakes (PTWI). We found that Cd concentrations were the limiting factor for mussels as a food. The weekly consumptions of 0.64–1.2 kg of fresh wild and 0.84–1.2 kg of fresh cultivated mussel would be sufficient to reach the PTWI_Cd, which may result in a risky weekly intake of Cd for long-term exposure. Moreover, weekly intake of 125 g mussels was used to calculate the dietary intake of trace elements by mussel consumption and compared with the prescribed PTWIs. Here, we found that there is no risk for human health for all investigated elements. In this case, the highest Cd level obtained in wild and in cultivated mussels represents 19.8 and 14.9% of the PTWI_Cd, respectively. This is the first study in Montenegro giving an assessment of the health risk from trace elements via the consumption of wild and cultivated M. galloprovincialis.