Seasonal variation in the balance between physiological mechanisms of feeding and digestion in Mytilus edulis (Bivalvia: Mollusca)

Seasonal variations in the balance between physiological mechanisms of feeding and digestion are considered among 45 to 57 mm shell length Mytilus edulis L. from southwest England. Mussels had been acclimated to standardised conditions of food availability in March, June and October, 1981. Results indicate that despite showing a negative relationship (P<0.001) with the efficiency of ¹⁵N absorption, coincident alterations of both ingestion rate (0.17 to 0.54 mg total Phaeodactylum tricornutum h^-1) and organic gut content (1.00 to 1.80 mg) represent significant mechanisms effecting the seasonal regulation of nutrient acquisition. Concurrent changes in absorption efficiency alone clearly exerted a relatively minor influence upon seasonal rates of absorption. Associated gut residence times of ¹⁵N-labelled material varied between 4 and 10 h. Furthermore, despite the ensuing deleterious effects, together with increased gut contents, absorption efficiencies among mussels that had been starved throughout acclimation were consistently maintained equivalent to those among individuals feeding normally. Such constancy was at least partially achieved by an increased residence time of material within diverticulae, relative to the digestive system as a whole. Finally, comparison of isotopic absorption efficiencies with those recorded for total organics and organic nitrogen has demonstrated differences of as much as 30% between gross and net uptake due, we suggest, to excretion of metabolically derived material within the alimentary canal.     

Labial palps of the blue mussel Mytilus edulis (Bivalvia: Mytilidae)

As a basis for understanding the functions of labial palps in the blue mussel Mytilus edulis, the structure and histology of palps were studied using light and scanning electron microscopy. Mussels used in the present study were collected in August 1993 and April 1994. The palp ridged surface is characterized by the presence of a smooth but densely ciliated dorsal fold, upon which rests the corresponding demibranch ventral region. The underside of the dorsal fold and the palp ridges fuse to form vestigial ciliated tracts. The dorsal fold is capable of contraction, allowing it to cover variable amounts of the ridged surface. Two different types of mucocyte are present on the palp ridged surface: subepithelial, glandular, acid-dominant secretion mucocytes and epithelial mucocytes characterized by neutral secretions. In histological section, these mucocytes appear to be concentrated on anatomical features known to intervene in particle handling. The anatomical and histological features of the smooth surface are typical of bivalve labial palps, except that the dense ciliation of the dorsal fold begins in the dorsal region of the smooth surface, indicating the possible origin of this feature. Previous studies on M. edulis point to the palps as the probable site for both ingestion volume control and particle selection; the anatomical basis of the present study should facilitate further research on these aspects.     

Digestive rhythms in the mussel Mytilus edulis

Mytilus edulis L., collected from a mid-tide level on the shore, showed rhythmic changes in mantle fluid pH, crystalline style pH, style length and total protein, and in the amylase activity in the digestive gland. These changes were correlated with the changes in tidal height. Style size may be related to extracellular digestion in the stomach. Style size and amylase content of the style were not significantly correlated with each other. The changes in amylase activity in the digestive gland confirmed the existence of a tidal rhythm for intracellular digestion in M. edulis.     

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

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

An ultrastructural study of Crassostrea angulata (Mollusca,Bivalvia) spermatogenesis

The cytoplasm of Crassostrea angulata (collected from the Atlantic coast of Northern Spain, Galiza, between 1992 and 1993) spermatogonia contains a few proacrosomal dense vesicles and intermitochondrial round granulo-fibrillar dense bodies. In primary spermatocytes, large cytoplasmic spaces appear occupied by large round and more scattered granulo-fibrillar dense bodies which are intermingled with cord-like dense structures and are in close association with mitochondria and the rough endoplasmic reticulum. As a result of Golgian activity, the number of dense proacrosomal vesicles increases and a flagellum arises from the centrioles. During the second meiotic division, several round dense bodies surrounding cord-like dense structures as well as dense proacrosomal vesicles, were observed between chromosomes. During spermiogenesis, no Golgian activity was noticed, and only the rough endoplasmic reticulum was observed in close association with proacrosomal vesicles. Similar results were observed for Ostrea edulis (collected from the Atlantic coast of southern Portugal, Algarve, between 1992 and 1993). Spermatozoa of both species are of the aquasperm type, and although their ultrastructure is quite similar, silver and phosphotungstic acid-chromic acid stainings enabled us to clearly distinguish both species at the ultrastructural level. The different morphological aspects of spermatogenesis in the Pteriomorphia and the use of silver and PTA stainings in the Mollusca are reviewed and discussed.     

Functional and environmental anaerobiosis in the razor-clam Ensis directus (Mollusca: Bivalvia)

In Ensis directus (Conrad) metabolic changes accompanying muscular activity and experimental anoxia were investigated. Octopine dehydrogenase, strombine dehydrogenase, alanopine dehydrogenase and lactic dehydrogenase were shown to be present in different parts of the body, but the activity of ODH always exceeded that of the other enzymes by far. Muscular activity connected with escape digging or evoked by electrical stimulation resulted in a decrease of stored phosphagen and in an accumulation of octopine. During recovery the normal concentrations of energy-rich phosphates were restored quickly, whereas octopine was removed more slowly. In experimental anoxia only minor quantities of L-and D-alanine, succinate, propionate, and acetate were formed. Quantitatively the most important endproduct was strombine, which rose linearly with the duration of anoxic incubation.     

Ice nucleation in the blue mussel (Mytilus edulis)

The ice nucleation temperatures of different homogenised organs of the mussel Mytilus edulis L. were examined. The stomach and the hemolymph had the highest nucleation temperatures. In the homogenised stomach the nucleation temperatures were fairly constant throughout the year, whereas the nucleation temperatures of the hemolymph increased in the cold season. Bacterial growth experiments, microfiltration, and experiments using antibiotics indicated that the nucleators were not of bacterial origin. The nucleation temperatures of natural seawater were approximately −9 °C, whereas seawater made from distilled water and sea salt had nucleation temperatures of about −17 °C. The nucleation temperatures of the seawater were reduced when the seawater was filtered by the mussels. However, no clear indication that the nucleators in the stomach were obtained from the seawater was found. Stomach homogenates from mussels kept in nucleator-free water had the same supercooling points as stomach homogenates from mussels kept in natural seawater. This indicates that the nucleators in the stomach are not obtained from the seawater. The temperature and light conditions examined in the present study did not significantly influence the hemolymph ice nucleation temperatures of mussels kept in the laboratory. Received: 24 October 1996 / Accepted: 16 November 1996     

Variability of the frequency of male neoteny in Ostrea puelchana (Mollusca: Bivalvia)

Ostrea puelchana (D'Orbigny), a larviparous oyster from the Atlantic coast of South America, comprises two types of individuals: (1) adults which live for several years and become sexually developed after one year, these show a rhythmic successive hermaphroditism with male, hermaphroditic and female phases; (2) neotenic individuals settled on the adults, which live for only about one year and function exclusively as males. Adult oysters were collected in February 1982 from a natural bed in the San Matias Gulf, Argentina, and were transferred to two sites 200 km to the south of their original habitat, where they were maintained under culture conditions. At the beginning of the study, 17% of adult oysters were observed bearing neotenic individuals. After two years, the number of adults in the female phase had decreased, and neotenic individuals had entirely disappeared. The presence of female-phase adults would seem to be necessary for the existence of neotenic males. Further analysis of the results revealed high adult density to be directly associated with greatly decreased numbers of adults in the female phase; this may play a role in the decrease of neotenic individuals.     

Trace metals in mussels (Mytilus edulis) from southwest Iceland

The temperate population of the tropical anemonefish Amphiprion clarkii has a seasonality of reproduction, larval settlement and growth. There was a considerable difference in size (15–47 mm) among 0-year olds in December, after the season of larval settlement and growth. This difference could be attributed mainly to a difference in the duration of the first growing season spent after settlement and a social inhibition of growth of the subordinate late settlers by frequent attacks by the dominant early settlers. The growth equation which was used to describe the growth of tagged individuals estimated that an anemonefish would get an opportunity to spawn within four of five years after settlement. Breeding adult pairs occupied almost all sea anemones essential to breeding within a 50×50 m study area, and all 17 new adults that matured from non-breeding juveniles to breeding adults between June 1983 and August 1985 were found in the territories from which either or both of the mated adults had disappeared. The new adults were large juveniles who had resided near those vacant territories before. These results suggest that the onset of breeding by an individual is not only determined by his age, but also by his ranking in the dominance hierarchy. Therefore, the difference in size among 0-year olds in December might give rise to the difference in age of the earliest breeding and might be a factor in making a female apply more reproductive effort at the beginning of the season.     

Freshwater Mussel,Lamellidens Marginalis (Lamarck) (Mollusca: Bivalvia: Unionidae) As an Indicator of River Pollution

Abstract

Analyses were made of heavy metals, manganese, nickel, copper, zinc and lead in water samples and soft body, shell and different tissues (gills, digestive glands, mantle and viscera) of the Unionid mussel, Lamellidens marginalis collected from two tributaries of the Cauvery river. Water samples from Station I contained higher concentrations of the metals than those from Station II. the concentration of metals in water at both stations were in the descending order: Mn > Zn > Pb > Ni > Cu. However, the concentrations of metals in the soft body were in the descending order: Zn > Mn > Pb > Ni > Cu at both stations in all size groups of mussels tested. the concentration of zinc maintained a linear relationship with the size of the mussels, but manganese showed a reverse trend. Small size (4-5 cm) mussels accumulated more manganese (105.5 μg.g^?1 dry wt.) than larger ones (7-8 cm; 6.5 μg.g^?1 dry wt.). Both young and old is accumulate the same level of lead, copper and nickel in the soft body. the order of concentrations of metals (Mn, Pb, Zn, Ni and Cu) in the shell of mussels from both stations coincided with the order of concentrations of background water except for lead. the accumulation of lead was higher in shell (30.4-36.2 μg.g^?1 dry wt.) than in soft body (6.4-12.0 μg.g^?1 dry wt.). the pattern of concentration of metals in the various tissues reveal that the digestive glands have greater ability than other tissues to concentrate most metals under study. the concentration factors for soft body, shell and different tissues are presented. the advantages in using the common mussel for biomonitoring of contaminants in water is also discussed.     

Le caryotype d'Aulacomya ater regia (Mollusca: Bivalvia: Mytilidae) des Iles Kerguelen

In 1983, specimens of the antarctic mussel Aulacomya ater regia Powell, 1957 were collected from the Kerguelen Islands and transported alive to the Zoological Station in Villefranche-sur-Mer, France. The chromosome complement of the mussel, as determined from branchial tissue, was found to consist of 13 pairs of chromosomes with terminal or subterminal centromeres. This species differs from the other species of Mytilidae studied up to the present in the number and the morphology of the chromosomes. A. ater regia is also unique among the known bivalves in displaying only terminal or subterminal centromeres.     

Peribuccal organs ofPlacopecten magellanicus andChlamys varia (Mollusca: Bivalvia): Structure,ultrastructure and implications for feeding

In order to better understand the structure of bivalve peribuccal organs and relate this to existing functional paradigms of their role in feeding, the labial palps of two scallop species,Placopecten magellanicus from the Bay of Fundy, Canada (1985 and 1986), andChlamys varia from the Bay of Brest, France (1986), were examined using histological techniques and electron microscopy. The ridged palp surface displays a uniformly dense ciliation with relatively few mucocytes; these are essentially concentrated in the region of the secondary ledge and may, through their secretory activity, determine the fate of particle masses in this area. The mucus secretions of the ridged palp surface are qualitatively different from those of the smooth palp surface. Mucocytes are much more abundant on the smooth palp surface, where it is suggested that their homogeneous secretions attenuate the potentially adverse effects of anteriorly-directed cleansing and swimming currents. Two other cell types are found in the palp epithelia: ciliated cells, which are very numerous on the ridged surface and relatively rare on the smooth surface, and non-ciliated epithelial cells, which are very numerous on the smooth surface and rare on the ridged surface, where they are confined to the palp margin. In addition to the mechanical role of the ciliated cells and mucocytes, the ultrastructural characteristics of the ciliated and non-ciliated epithelial cells indicate a dichotomy of function between the ridged and smooth surfaces. The ridged surface epithelial cells present an ultrastructural specialization in the absorption of dissolved and colloidal matter, suggesting an accessory nutritive role, whereas the smooth surface simple epithelial cells show signs of active molecular synthesis. No specialised sensory cells were observed on the ridged surface; it is therefore not yet possible to conclude whether the labial palps are capable of selection based on individual particle characteristics.     

Temperature-time relationships for survival of embryos and larvae of Mulinia lateralis (Mollusca: Bivalvia)

The influence of temperature over 8 time intervals on survival of cleavage stages, trochophore larvae, and straighthinge veliger larvae of Mulinia lateralis (Say) was investigated using a thermal gradient apparatus. There was a direct relationship of mortality with increased period of exposure. As the clams aged, temperature tolerance increased, with cleavage stages being most sensitive to higher temperatures and straight-hinge larvae least sensitive. Multiple-regression equations were developed to allow prediction of percentage mortalities under different conditions of temperature and time exposure. Entrainment of M. lateralis embryos and larvae in the cooling-water systems of steam-electric power plants should be as short as possible to keep mortality toa minimum.Contribution No. 550 of the Natural Resources Institute, University of Maryland, and Contribution No. 531 of the Virginia Institute of Marine Science.     

Advanced age for sexual maturity in the ocean quahog Arctica islandica (Mollusca: Bivalvia)

A study of gonadal state in 39 small specimens of Arctica islandica Linné collected in April 1976 from the Middle Atlantic shelf of North America suggests that sexual maturity is reached at a later age than has been reported for other shelf bivalves. Eight of the specimens had gonads in the undifferentiated state. On the basis of annual internal growth banding in the shells of these immature specimens, age ranged between 4 and 14 years. Average age was 9.38 years (standard deviation, s,= 3.54). Shell length ranged from 24 to 47 mm (average 38.85 mm, s=9.25). Differentiated gonads were found in specimens as young as 6 years. Age of maturity shows a wide range and may be dependent upon growth rate and locality.     

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     

Seasonal variation in mineralization rates (C-N-P-Si) of mussel Mytilus edulis biodeposits

To determine seasonal variability in mineralization dynamics of mussel biodeposits, we applied a multiple-element approach measuring mineralization rates of carbon (C), nitrogen (N), phosphorus (P) and silicate (Si) during three periods (March, August and November). The results of this study showed that mineralization rates vary between seasons and between elements and that mineralization dynamics were influenced by both temperature and biodeposit nutrient composition. Mineralization rates were 3.2 ± 0.4 mmol C, 0.17 ± 0.04 mmol N, 0.06 ± 0.02 mmol P and 3.91 ± 3.75 mmol Si per gram biodeposit (DW) per day, which represented 24 % of the particulate organic C and 17 % of the particulate organic N in mussel biodeposits. Seasonal variability was largest for Si mineralization with 60–80-fold higher rates measured in March compared to August and November. This difference is most likely related to the difference in biodeposit nutrient composition. It was furthermore shown that the labile fraction of biodeposits became mineralized after, respectively, 18, 9 and 13 days during the experimental periods in March, August and November. This indicates that temperature enhances biodeposit decomposition with approximately 2–3 times faster turnover at a 10 °C temperature interval (Q ₁₀ ).     

First evidence of gender-related differences in immune parameters of the clam Ruditapes philippinarum (Mollusca, Bivalvia)

For the first time, gender-related differences in some important functional parameters of haemocytes from the clam Ruditapes philippinarum have been studied. Clams were collected during the pre-spawning phase, when sex can be distinguished by microscopic observation of gonadal smears, in order to ascertain whether the two sexes reach the stressful spawning period with different degrees of immunosurveillance. Total haemocyte count (THC), haemocyte size frequency distribution, capability of haemocytes to assume the vital dye neutral red (NR, indicative of endocytotic activity), lysozyme-like and acid phosphatase activities in both haemocyte lysate (HL) and cell-free haemolymph (CFH), and superoxide dismutase (SOD) and catalase (CAT) activities in HL were all evaluated. No differences in THC values were found between the two sexes, but differing haemocyte size frequency distributions were observed: the fraction of larger haemocytes (7–9 μm diameter, 200 fl volume) was markedly increased in females and that of smaller haemocytes (<5 μm diameter, <200 fl volume) markedly increased in males. Interestingly, when haemocytes were observed under the light microscope, a significantly higher percentage of granulocytes was found in haemolymph from females, compared with that of males. Significantly increased NR uptake was recorded in female haemocytes. No significant variation in lysozyme-like activity was observed in HL, whereas in CFH, enzyme activity was significantly higher in females. HL acid phosphatase activity was significantly higher in males with respect to females; no significant variations in enzyme activity were observed in CFH. HL SOD and CAT activities were significantly higher in females than in males. Overall, on the basis of the cell parameters studied here, gender-related differences in immune parameters were found in R. philippinarum, indicating that females have more active haemocytes than males during the pre-spawning period.     

Gill function and particle transport in Placopecten magellanicus (Mollusca: Bivalvia) as revealed using video endoscopy

The technique of endoscopic video observation was used to study feeding processes of Placopecten magellanicus (Gmelin), collected from Bull Arm, Newfoundland in August 1991 and 1992, under near-natural feeding conditions. The fate of captured particles depended on the extent of ingestive or handling capacity saturation. Under low (1 to 10 particles l^-1) to medium (10 to 20 particles l^-1) particle concentrations, most particles were incorporated in continuous anteriorly directed slurries in the dorsal ciliated tracts of the gill arch and dorsal bends. As particle concentration or exposure time to the lower particle concentrations increased, four endogenous mechanisms of ingestion volume control were increasingly observed: (1) rejection of dense mucus-particle masses from the principal filament troughs onto the ventrally beating cilia and associated currents of the ordinary filament plicae, counter to and below the incoming pallial current maintained by the principal filament cilia; (2) intermittent stopping of the anteriorward flow in the dorsal ciliated tracts; (3) reduction or cessation of input from the principal filaments to the dorsal ciliated tracts; (4) detachment of the dorsal bends from the mantle to establish a shunt from the infrabranchial to the suprabranchial cavity. Chemical and histochemical tests of purified fluid withdrawn from the dorsal ciliated tracts indicate that mucus is present at all particle concentrations. Mucus therefore participates both in normal feeding and in ingestion volume control on the bivalve gill, although different mechanisms, and types of mucus, effect transport of material in the dorsal (feeding) and ventral (cleaning) ciliated tracts.     

Burrow morphology,biometry, age and growth of piddocks (Mollusca: Bivalvia: Pholadidae) on the south coast of England

Biometry and growth of three piddock species Pholas dactylus, Barnea candida and B. parva, from chalk and clay substrata were investigated between 1999 and 2000 at five low shore locations along the south coast of England. Piddock burrow shape was significantly different (P<0.01) amongst the locations. Burrows at Lyme Regis showed the largest (height/maximum diameter) ratio (4.86±2.00) whilst those at Compton the lowest (3.73±1.62). Using the method of Bhattacharya, the population structure of P. dactylus, B. candida and B. parva was separated into eight, three and five modal size classes, respectively. Age and shell growth were determined from the number and spacing, respectively, of annual growth lines present in acetate peel replicas of shell sections. The von Bertalanffy growth (VBG) equation fitted the size at age data obtained for P. dactylus and B. candida (L and K were 79.3±13.8 mm and 0.0011±0.22 and 29.6±1.5 mm and 1.17±0.47, respectively) whilst the size at age data for B. parva were linearly related and did not fit the assumptions of the VBG equation. Male and female gonads were mature and piddocks competent to spawn between June and September 1999, with settlement of juveniles observed between November 1999 and February 2000. A significant relationship between burrow aperture diameter and age of the occupant piddock was established for the three species. Burrow morphology and spatial distribution of burrows were influenced by substratum hardness and population density. Based on estimates at Lyme Regis, piddocks are capable of removing up to 41% of the shore substratum to a depth of 85 mm over their lifespan (12 years), significantly compromising the structural stability of the soft rock shores they inhabit and contributing to bioerosion.Communicated by O. Kinne, Oldendorf/Luhe