Endogenous circatidal rhythm in the Manila clam Ruditapes philippinarum (Bivalvia: Veneridae)

Manila clams, Ruditapes philippinarum, removed from their natural environment and maintained for 9 weeks in continuously immersed conditions exhibited a clear endogenous circatidal rhythm in oxygen consumption. The clams exhibited a semidiurnal rhythmicity in oxygen consumption after showing a diurnal pattern in the first few days (5 to 7 d) of the experiment. The results of the present study indicate that activity rhythms of clams are controlled not only by exogenous factors, but also by an endogenous circatidal periodicity. Received: 8 April 1998 / Accepted: 1 February 1999     

Effects of salinity on endogenous rhythm of the Manila clam, Ruditapes philippinarum (Bivalvia: Veneridae)

The Manila clam Ruditapes philippinarum, an intertidal bivalve, was exposed to different salinity regimes (from 31.0–31.7‰ down to 20‰, 15‰, 10‰, 5‰), and the endogenous rhythm in its oxygen consumption was studied using an automatic intermittent-flow respirometer. When exposed to salinities reduced from 31.5‰ to 20‰ and 15‰ under otherwise constant conditions, the clams recovered a clear endogenous circatidal rhythm in their oxygen-consumption rate after having dampened periods of 12 h and 48 h, respectively. At salinities less than 10‰, however, the oxygen-consumption rate was depressed greatly at the beginning of the experiment for about 36 h and then increased to a level higher than normal, but the rhythm of oxygen consumption was not recovered. The results of this study indicate that the Manila clam, a euryhaline organism, cannot maintain a normal metabolic activity at a salinity lower than 15‰. All clams were dead after exposure at a salinity of 5‰ for 7 days. Received: 28 February 2000 / Accepted: 26 August 2000     

Acid and alkaline phosphatase activities in the clam Ruditapes philippinarum

Acid and alkaline phosphatase activities have been partially characterized in Ruditapes philippinarum (Adams and Reeve, 1850). Two activity peaks at pH=4.5 and pH 10.5 were detected in the gill, digestive gland, mantle, siphon and foot. Acid phosphatase activity was higher than that of alkaline phosphatase. The highest activity for both enzymes was observed in the digestive gland and, in decreasing order, the gill, foot, siphon and mantle. Alkaline phosphatase activity was similar in the mantle, siphon and foot. K _m values were determined for both enzymes in the gill and digestive gland. Hill coefficients were near 1, indicating no allosteric behaviour for either enzyme in the two organs. The optimum temperature was the same for acid phosphatase in both gill and digestive gland (50 °C), while for alkaline phosphatase it differed for these two organs (gill, 40 °C; digestive gland, 35 °C). The apparent activation energy was obtained from Arrhenius plots, and ranged from 8.61 kcal/mol for alkaline phosphatase in the gill, to 10.84 kcal/mol for acid phosphatase in the digestive gland. The effects of metals (1 mM conc) on both enzyme activities were assayed in vitro. Hg strongly inhibited the enzyme activities in the gill and digestive gland, probably because of its affinity for the sulphydryl group. Histochemically, acid phosphatase in the gill was located in a granular form throughout the gill cells, but was undetectable in the ciliate epithelium of the gill filaments. Alkaline phosphatase was located in the gill skeleton. Clam size and phosphatase activities were inversely related, probably reflecting a decrease in shell deposition with inereasing size. As a function of season, both enzymes were present in lowest amounts in winter, when undifferentiated sex cells were predominant in the germinative epithelium, and highest in summer, when ripe individuals of both sexes were more frequent.     

A system dynamics model for the management of the Manila clam, Ruditapes philippinarum (Adams and Reeve, 1850) in the Bay of Arcachon (France)

The Manila clam Ruditapes philippinarum (Adams and Reeve, 1850) is one of the mollusc species that, driven mainly by the shellfish market industry, has extended throughout the world, far beyond the limits of its original habitat. The Manila clam was introduced into France for aquaculture purposes, between 1972 and 1975. In France, this venerid culture became increasingly widespread and, since 1988, this species has colonised most of the embayments along the French Atlantic coast. In 2004, this development resulted in a fishery of ca. 520 t in Arcachon Bay.     

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.     

Preliminary data on PAH metabolism in the marine mussel Mytilus galloprovincialis from Arcachon Bay,France

Oxidation of polycyclic aromatic hydrocarbons (PAH) and epoxide-conjugation capacities were examined from mussels, Mytilus galloprovincialis, collected from a clean site at the entrance to Arcachon Bay (south-west France) in 1984, using subcellular preparations from whole bodies and digestive glands. The existence of low-cytochrome P-450-dependent oxygenase activities [cytochrome P-450 content, NADPH cytochrome c reductase and benzopyrene monooxygenase (BaPMO) activities] was confirmed, as well as the presence of a comparatively important capacity for epoxide conjugation [epoxide hydrolase (EH), glutathione S epoxide-transferase (GST) activities and total cytosolic-glutathione content]. The optimal incubation temperatures for the individual in vitro enzyme activities in the mussel (25°C for BaPMO, 31°C for EH and GST) were lower than that required for optimal mammalian enzyme activities. Except for cytochrome P-450 content, there was no significant difference in enzyme activities recorded in whole-body and digestive gland fractions.     

Interactions between two introduced species: Zostera japonica (dwarf eelgrass) facilitates itself and reduces condition of Ruditapes philippinarum (Manila clam) on intertidal flats

Dwarf eelgrass (duckgrass; Zostera japonica) and Manila clams (Ruditapes philippinarum) are two introduced species that co-occur on intertidal flats of the northeast Pacific. Through factorial manipulation of clam (0, 62.5, 125 clams m⁻²) and eelgrass density (present, removed by hand, harrowed), we examined intra- and interspecific effects on performance, as well as modification of the physical environment. The presence of eelgrass reduced water flow by up to 40% and was also observed to retain water at low tide, which may ameliorate desiccation and explain why eelgrass grew faster in the presence of conspecifics (positive feedback). Although shell growth of small (20–50 mm) clams was not consistently affected by either treatment in this 2-month experiment, clam condition improved when eelgrass was removed. Reciprocally, clams at aquaculture densities had no effect on eelgrass growth, clam growth and condition, or porewater nutrients. Overall, only Z. japonica demonstrated strong population-level interactions. Interspecific results support an emerging paradigm that invasive marine ecosystem engineers often negatively affect infauna. Positive feedbacks for Z. japonica may characterize its intraspecific effects particularly at the stressful intertidal elevation of this study (+1 m above mean lower low water).     

Accumulation of lead and zinc and effects on activities of catalase and GPT in the clam Ruditapes philippinarum A

Acute toxicities of Pb and Zn to clam (if. philippinarum) were determined by a semistatic bioassay. The 96 h LC50 values were 14.28 mg/L for Pb, and 16.40 mg/L for Zn. Exposure of R. philippinarum to various concentration of lead and zinc for a period of 4 days showed that different organs have different capacities for accumulating metals, with high levels in the gill, followed by soft body, and lower amounts in the cavity fluid. Accumulation tests showed that R. philippinarum is a strong accumulator of Pb and weaker accumulator of Zn. The effect of lead on activities of Catalase and GPT were studied in soft body of R. philippinarum. The result showed that the activities of CAT in soft body decreased at all exposure groups, the activities of CAT were negatively correlated with exposure concentration of lead, the special inhibition effect was observed in exposure group of Zn, the CAT activities significantly decreased at lower Zn concentration, then increased slowly with the Zn concentration, finally, decreased to the lowest. The activities of GPT in soft body were significantly increased at high exposure concentrations.     

Stable isotopes changes in the adductor muscle of diseased bivalve Ruditapes philippinarum

In this article, we show how a disease could bias stable isotope analyzes of trophic networks and propose a strategy in the choice of tissues to be analyzed. In the past few years, a new pathology (brown muscle disease or BMD) affecting the posterior adductor muscle of Ruditapes philippinarum has emerged in Arcachon Bay. BMD induces a necrosis of muscle tissues which become infused by conchiolin and hence calcified. As muscle of mollusks are often used for trophic food webs studies through stable isotopic analyzes, this work investigated the effect of BMD on carbon and nitrogen isotopic ratios of anterior and posterior adductor muscles of clams collected in February and August 2007. Infected clams displayed a lower condition index and a posterior adductor muscle δ¹³C enrichment of 1.2‰ in February and 0.7‰ in August. δ¹⁵N of posterior muscles was however not affected by the disease. Anterior muscle of diseased clams remained healthy and displayed the same isotopic signature as both posterior and anterior muscular tissues of healthy clam. Acidification significantly depleted δ¹³C in posterior muscles of infected clams, suggesting calcification, contrary to anterior muscles of infected clam and to both muscles of healthy clams, where no effect was observed. An X-ray diffractometry analysis confirmed the presence of CaCO₃ (aragonite). Trophic food web studies relying on stable isotope ratios should utilize only healthy animals or anterior adductor muscles when expertise in mollusk pathology is lacking.     

Potential of bivalves' secondary settlement differs with species: a comparison between cockle (Cerastoderma edule) and clam (Ruditapes philippinarum) juvenile resuspension

Juvenile bivalves may be dispersed by entering a bysso-pelagic phase where they drift through the water mass aided by a long thread. The ability to resuspend and control the specific weight in two bivalve species, the cockle Cerastoderma edule (L.) and the Japanese clam Ruditapes philippinarum (Adams and Reeves), was documented with juveniles through flume and still-water experiments. Cockle juveniles initially placed on an unsuitable substratum were exposed to two shear velocities (u _*). At the end of the experiment, 42 (±15)% (for u _*=0.51 cm s⁻¹) and 79 (±9)% (for u _*=0.99 cm s⁻¹) of individuals were retrieved from the sand area which represents only 8% of the total flume surface. Most juveniles (70.5%) with shell lengths <2.5 mm migrated from the unsuitable Plexiglas substratum to the sand array by resuspension in the water column. The percentage was lower (21.5%) for larger individuals. The same experimental design was applied to clams, which immediately adhered to the Plexiglas substratum and remained attach to it. Sinking rates of live and dead specimens of both species were measured in a 1 m long transparent PVC tube. Cockle fall velocities showed severe deceleration, probably due to byssus secretion (up to 15-fold slower than dead cockles), sometimes interrupted by brutal acceleration probably due to byssal rupture. Cockles were able to reduce their sinking rate for shell lengths up to 4.25 mm. By contrast, clam sinking rates were constant, and similar to dead clam sinking rates. Specific weights of all experimental juveniles were calculated in relation to their lengths, and their passive motion into the boundary layer was theoretically assessed with Shields curve. In short, C. edule and R. philippinarum can both exhibit dense populations in the field with a good capacity to colonize, although juveniles display different abilities to resuspend in the water column. Received: 27 January 1997 / Accepted: 13 February 1997     

Genetic population structure of the recently introduced Asian clam,Potamocorbula amurensis,in San Francisco Bay

The genetic population structure of the recently introduced Asian clam, Potamocorbula amurensis, in San Francisco Bay was described using starch gel electrophoresis at eight presumptive loci. Specimens were taken from five environmentally distinct sites located throughout the bay. The population maintains a high degree of genetic variation, with a mean heterozygosity of 0.295, a mean polymorphism of 0.75, and an average of 3.70 alleles per locus. The population is genetically homogeneous, as evidenced from genetic distance values and F-statistics. However, heterogeneity of populations was indicated from a contingency chi-square test. Significant deviations from Hardy-Weinberg equilibrium and heterozygote deficiencies were found at the Lap-1 locus for all populations and at the Lap-2 locus for a single population. High levels of variability could represent a universal characteristic of invading species, the levels of variability in the source population(s), and/or the dynamics of the introduction. Lack of differentiation between subpopulations may be due to the immaturity of the San Francisco Bay population, the general purpose phenotype genetic strategy of the species, high rates of gene flow in the population, and/or the selective neutrality of the loci investigated.     

Selective feeding in shellfish: size-dependent rejection of large particles within pseudofaeces from Mytilus edulis, Ruditapes philippinarum and Tapes decussatus

Mytilus edulis L., Ruditapes philippinarum (Adams & Reeve) and Tapes decussatus L. were fed particles of the same shape (spherical), the same density (2.1 g cm⁻³) and the same chemical composition (SiO₂), but which varied in diameter from 5 to 37 μm. Findings obtained at different particle concentrations (mean ± SD) of 51 ± 2, 105 ± 18 and 171 ± 17 mg l⁻¹ invariably indicate that significant proportions of all particles with diameters larger than from between 7.5 and 22.5 μm were preferentially rejected as pseudofaeces. We define the preferential ingestion index (PII) as the ratio between average particle volume in pseudofaeces and average particle volume in food. Whatever the particle concentration or the species, this PII was always statistically higher than 1. Irrespective of particle concentration, PII values in M. edulis were lower than in T. decussatus (averages of 1.2 and 2, respectively). PII values in M. edulis were also lower than in R. philippinarum maintained at particle concentrations above 171 ± 17 mg l⁻¹. We suggest that preferential size-dependent rejection of larger particles could be of significant adaptive value in the natural environment, either if there are large inorganic particles, or if the average organic content of smaller particles is higher. Received: 11 January 1997 / Accepted: 8 March 1997     

Linear alkylbenzene sulphonates (LAS) and bioaccumulation of heavy metals (Cu and Pb) in Ruditapes philippinarum

The effect of linear alkylbenzene sulphonate (LAS) on copper and lead bioaccumulation in Ruditapes philippinarum was studied. The in vitro and in vivo effect of LAS on acid and alkaline phosphatase activities was also examined. For lead, significant differences (P<0.01) were observed between organisms exposed to lead and to the mixture of LAS and lead. In the case of copper, these differences were only significant in the digestive gland (P<0.05). In vitro acid phosphatase was inhibited by LAS and the mixture of LAS with copper showed the effect of decreasing the enzymatic activity. Nevertheless, in vivo no significant differences in the digestive gland and gills (P < 0.05) were observed.     

Reproduction of the invasive slipper limpet, Crepidula fornicata, in the Bay of Brest, France

The reproduction of Crepidula fornicata was studied in the Bay of Brest in order to characterise the first step of the reproductive cycle of this invasive species. The survey was carried out from 2000 to 2003 and different parameters were measured, namely, the percentage of the different sexual stages, the straight length of the shell and the percentage of brooding females using a survey of the embryonic development and the fecundity. The juvenile frequency increases generally from mid-June or mid-August, depending on the year. In 2001 and 2003, a first peak was observed as early as May, but it was followed by a rapid disappearance of the individuals. The sex-ratio female/male increased from 0.22 to 0.46 between 2001 and 2003. The sex change between intermediates and females took place mainly in summer and was well marked in 2001 and 2003. The survey of the embryonic development in the egg capsules brooded by the females provided an annual phenology of the laying and hatching processes. The laying period extends from February to September with three to four major periods of egg-laying per year and corresponding hatching periods about 1 month later. Each female lays two to four times per year on average. The first egg-laying concerned fewer females than subsequent ones, except in 2003, and exhibited a higher fecundity. The annual mean of the number of eggs for each stage was not significantly different, thus indicating no significant mortality rate during embryonic development. For the C. fornicata population in the Bay of Brest, several reproductive characteristics tend to highlight its invasive capacity: (1) a long reproductive period, (2) reproduction in a ‘multi-trials’ process equivalent to a spreading out of the risks and (3) a relatively high fecundity.     

Growth,mortality and recruitment of the yellow clam Mesodesma mactroides on Uruguayan beaches

We studied some aspects of the population dynamics of the yellow clam Mesodesma mactroides (Deshayes, 1854) which inhabits the eastern sandy beaches of Uruguay. Observations were made from March 1983 through March 1985 (harvesting season), and additional data from January 1988 to December 1989 (closed season) were also included in the analyses. Parameters of growth and mortality were estimated by methods that make use of age or length information. An age/length relationship was constructed by counting growth rings. Results were consistent with those obtained with modal class progression analysis and other length/frequency methods. The growth curve obtained from the age/length relationship did not account for seasonality in growth rate, and a seasonal growth model explained growth oscillations in a better way. The instantaneous coefficient of natural mortality (M) estimated for the closed season was 1.64, whereas an increase in fishing mortality (F) was observed during 1984. Age composition did not change during the 2 yr of study, and the first 6 mm class accounted for over 50% of the population. The main contribution of recruits was observed between late summer and early fall (February to April), and the recruitment pattern suggested one main settlement period per year. Growth and recruitment success appeared to be regulated by density-dependence processes. Some implications for management are considered in this context. Methodological aspects concerning the reliability of empirical equations and programs for estimating growth and mortality are discussed.     

Growth and reproduction in the mesopelagic-boundary squid Abralia trigonura

Abralia trigonura is a small squid that is an important member of the mesopelagic boundary community around Hawaii. Squids were aged by examining statolith growth increments, and various reproductive parameters were measured. Female longevity is 6 mo, and they become sexually mature at 3.5 mo. Male longevity is the same, but they become sexually mature at 2.5 mo. This species appears to be a multiple spawner, and peculiarities in the number of eggs found in the oviducts suggests that they spawn every few days. Possible advantages of this reproductive mode are examined.     

Variability of biochemical composition and size distributions of seston in the euphotic zone of the Bay of Biscay: implications for microplankton trophic structure

Particle-size distributions and several biochemical components of seston were studied from October 1986 to December 1987 in surface waters of the Bay of Biscay. Variance partitioning of hydrographic and seston parameters indicates that, although seasonal variability related to the thermal cycle was important, most of the differences in seston concentration were caused by spatial components of variance. Changes in the vertical structure of the water column appeared to be the principal source of variability. Phytoplankton blooms in spring and fall and the effects of the coastal upwelling and the thermocline during summer were traced using seston concentration and biochemical composition. The different seston measurements were scaled according to their correlations with total concentration and particle size. Photosynthetic pigment biomass was related to both large and small partieles. However, concentrations of particulate proteins, lipids and carbohydrates were more related to small particles. Although not specifically analysed, low concentrations of inorganic particles and detritus can be expected in the study area, thus most of the particulate organic matter was associated with small, non-pigmented organisms, particularly in surface waters during the period of thermal stratification. The variability in concentrations of total seston and particulate organic matter was mainly due to variations in particulate proteins, lipids and carbohydrates; pigment concentrations were of secondary importance. Expressed as the ratio chlorophyll a: particulate protein-nitrogen, phytoplankton constituted an important fraction of the microplankton biomass only during spring blooms, when it averaged 75% of the particulate protein-nitrogen. In constrast, <30% of protein-nitrogen was related to chlorophyll a during the summer. These results suggest that an oligotrophic structure based on the microbial-loop prevails in microplankton assemblages of surface waters for most of the year.     

Growth,reproduction and nutrition of some Barents Sea hydromedusae in natural aggregations

Mass aggregations of the hydromedusae Tiaropsis multicirrata and Rathkea octopunctata have been studied in the Jarnyshnaya Fjord on the Kola Penninsula during the period of domination of the summer neritic plankton complex. The mass aggregations persist at least over periods of 2 months. No hyrometereological parameters could be established that would conceivably favour such aggregations. The factor most likely responsible for creating the aggregations is assumed to be related to the stable level of the summer zooplankton, production in the study area. The vast individual numbers of hydromedusae as well as their polyphageous feeding behaviour and high growth rates suggest that the medusae represent the most substantial consumers of plankton in inshore waters. The low and highly variable percent of medusae individuals which reveal food in their intestinal system lead the authors to assume additional food sources: nannoplankton, bacteria, and dissolved organic matter.     

Seasonal change of species composition,abundance, and reproduction of solitary entoprocts in Okinawa Island,the Ryukyu Archipelago,Japan

Monthly samplings of solitary entoprocts were carried out at Mizugama and Ginowan Fishery Port, Okinawa Island, the central Ryukyus, Japan from November 2000 to November 2001. We examined the entoproct populations on the glass slides that had been placed at the two sites for 2 months, and a total of 18,322 individuals of solitary entoprocts were recorded. They were Loxomitra mizugamaensis, L. ryukyuensis, L. tetraorganon, Loxosomella aloxiata, L. intragemmata, L. lecythifera, L. monocera, L. stomatophora, and Loxocorone allax. Our analyses indicate that (1) the abundance of solitary entoprocts in this region is high in summer and low in winter, (2) most species reproduce by budding all year round, and (3) sexual reproduction is restricted to a short period in summer (July or August).