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Micronuclei (MN), nuclear buds (NB), fragmented-apoptotic (FA) and bi-nucleated (BN)cells were analysed in gills of bivalve mollusks Macoma balthica and Mytilus edulis collected from five study locations in Lithuanian territorial waters of the Baltic Sea. The frequency of micronuclei varied from 1.28 to 3.63‰ (MN/1000 cells), of nuclear buds — from 0.50 to 1.49‰, fragmented-apoptotic cells from 0.53 to 1.72‰ and of bi-nucleated cells from 1.51 to 2.23‰. The highest level of MN and bi-nucleated cells was determined in mollusks from the offshore Station 65, nuclear buds and fragmented-apoptotic cells from Station 1B located close to the Butinge oil terminal. A comparatively high frequency of MN was observed in bivalves from the Butinge oil terminal area - 3.38‰ in Station 1B, 2.85‰ in Station B06 in June and in mollusks from Station N-4 (2.66‰) located nearby the oil platform D-6. [ABSTRACT FROM AUTHOR]

Abstract: The aims of the study were to determine the contents of glycogen, organic carbon and nitrogen in the Baltic clam Macoma balthica tissue and to find out whether there is any difference in fitness and chemical composition between individuals living in sediments with different pore-water hydrogen sulphide concentrations. It was found that hydrogen sulphide permanently present in the sediment at concentrations exceeding 513μmoldm−3 at the middle depths of the Gulf of Gdańsk (60m) had no adverse influence on the fitness of M. balthica and did not reduce the organic compound content in its tissue. This is very probably because of the presence of oxygen in the near-bottom water. Interestingly, animals living in such concentrations of hydrogen sulphide had heavier shells and a greater soft tissue weight. Moreover, the contents of organic matter and glycogen in their soft tissues were significantly higher. Glycogen was 30% higher on average, ranging from 11.1 to 30.3% AFDW during the year. At all the stations during the year, organic carbon in M. balthica tissue varied from 50.3 to 55.2% AFDW, and organic nitrogen from 7.7 to 11.7% AFDW. [Copyright &y& Elsevier]

Metal concentrations of Cu, Fe, Mn, Ni, Pb and Zn in an infaunal facultative deposit-feeding bivalve, the Baltic clam Macoma balthica, in the Gulf of Gdansk (southern Baltic Sea) were assessed and compared to selected concentrations of metals in the environment. Between October 1996 and September 1997, dissolved and easy extractable (by 1M HCl) metal fractions of total suspended particulate matter (TPM) in the overlying water and of surficial sediments (<63μm) were measured monthly at five sublittoral sites in the Gulf of Gdansk, and accumulated tissue metal concentrations in M. Balthica were determined simultaneously. The study highlights the importance of sediment geochemistry as a factor modifying ambient trace metal bioavailabilities. Surficial sediments appeared to contribute most to the accumulation of Cu and Pb in M. Balthica, reflecting the high metal availability in the Gulf. Assimilation of Cu from sediments is controlled by Mn components possibly through an inhibitory effect of Mn oxyhydroxides, while Pb accumulation from sediments depends on the organic content of the sediment. A dual metal uptake pathway, with a suspended particulate-bound fraction and surficial sediments, was apparent for Mn and Zn. Partitioning of Mn in sediments was related to the concentration of labile Fe, with increased levels of Fe tending to inhibit the accumulation of Mn by the clam. Tissue accumulated Zn might have been altered by the clam’s internal regulation, making Zn tissue concentrations, to some degree, independent of its environmental level. The principal source of Ni accumulated by the clams exists in the soluble phase. [Copyright &y& Elsevier]

Abstract: Ecosystem engineers can influence community and ecosystem dynamics by controlling resources, modifying the flow of energy or biomass, or changing physical characteristics of the habitat. Invasive hybrid cordgrass (Spartina alterniflora×Spartina foliosa) is an ecosystem engineer in salt marshes in San Francisco Bay, California, U.S.A. that raises intertidal elevations and may be either increasing C 4 plant carbon input into food webs or tying up carbon in a form that is not usable by consumers. A manipulative experiment compared abundance, growth, and stable isotope (δ13C and δ15N) composition of the clam Macoma petalum (=M. balthica) among native marsh, hybrid Spartina, and mudflats in central San Francisco Bay. We found higher densities (individuals m−2) of M. petalum on mudflats compared to either native or hybrid Spartina (p <0.001). Macoma petalum shell growth was significantly greater in mudflats than in either vegetation type in 2002 (p =0.005) but not 2003. Differences in shell growth between native and hybrid Spartina were not significant. Stable isotope results showed differences between habitats in δ13C but not δ15N. Carbon signatures of M. petalum placed in Spartina were much more depleted than the isotopic signature of Spartina. Neither native nor hybrid Spartina appears to be a significant carbon source for M. petalum in San Francisco Bay, and we found no evidence that hybrid Spartina contributes carbon to M. petalum beyond what is provided by S. foliosa, despite the hybrid''s much greater biomass. Our results show that loss of mudflat habitat, rather than increased input of C 4 carbon, is the greatest effect of the invasion of hybrid Spartina on M. petalum. [Copyright &y& Elsevier]

Abstract: The energy flow in various bivalve species (Macoma balthica, Mya arenaria, Cerastoderma edule, Mytilus edulis and Crassostrea gigas) in different habitats (intertidal, subtidal and offshore areas) of Dutch coastal waters was analysed by comparing growth observations in the field with model simulations by means of the Dynamic Energy Budget (DEB) theory. The reconstruction of the seasonal food conditions suggested that with respect to food consumption, M. balthica and C. edule are the most successful species, followed by M. arenaria, and M. edulis and C. gigas. Positive growth was correlated with the beginning of the spring bloom in primary production and food limitation between species was lowest during this period. After the spring bloom, severe food limitation during the summer period was found. As a result, negative body growth occurred during most of year. DEB model simulations both on an annual and a seasonal basis also showed that, at the prevailing water temperatures, growth of the different species was not maximal. It remains uncertain whether the suboptimal food conditions were the consequence of food limitation or also of reduced filtration efficiency due to the high sediment load in the water. [Copyright &y& Elsevier]

Abstract: Dynamic energy budgets are used for the description of the energy flow through individual organisms from the assimilation of food to the utilisation for maintenance, growth, development and reproduction. In this paper, a procedure for estimation of the parameters of Kooijman''s Dynamic Energy Budget (DEB) model is introduced and subsequently parameters are estimated for the following Northeast Atlantic bivalve species: the Baltic clam Macoma balthica (L.), the sandgaper Mya arenaria L., the cockle Cerastoderma edule (L.), the blue mussel Mytilus edulis L. and the Pacifc oyster Crassostrea gigas (Thunberg, 1793). For none of the species, a complete set of parameters could be compiled. A special protocol was developed to account for missing values and to achieve consistency between parameters. Species were similar in their optimal temperature range, as reflected in a common Arrhenius temperature of 5800 K, which corresponds with a Q10 of 2. Differences between species were observed in width of the optimal temperature range. The taxonomic relatedness between species was reflected in similar volume-specific maintenance costs, costs for growth and almost similar maximum storage density of energy. Species differed in their maximum surface area-specific assimilation rate by a factor of 6 and in the fraction of energy allocated to reproduction (ranging from 0.15 to 0.50). These differences are reflected in the maximum theoretical total shell length of the species, which varied from about 3 cm in M. balthica, 6 cm in C. edule, 15 cm in M. arenaria and M. edulis and 45 cm in C. gigas. [Copyright &y& Elsevier]

Abstract: The objective of the DEBIB project (Dynamic Energy Budgets in Bivalves) was to combine the French expertise of bivalve ecophysiology and modelling with the Dutch expertise in ecology and modelling to achieve the development of a general framework based on Dynamic Energy Budgets [Kooijman, S.A.L.M., 2000. Dynamic Energy and Mass Budgets in Biological Systems. Cambridge University Press, Cambridge] for the analysis of the functioning of bivalve species (growth, reproduction) in relation to the environmental conditions (water temperature, food conditions). The collaboration has yielded promising insights for studying life strategies of different bivalve species sharing the same resources and new perspectives (i) for coupling ecophysiological models with population dynamics and ecosystem approaches and (ii) for addressing important issues such as carrying capacity, trophic interactions between bivalves and sustainability of shellfish resources. Fields for future research include improvement of parameter estimation procedures and analysis of intraspecies variability in parameters. [Copyright &y& Elsevier]

Abstract: In the Baltic clam Macoma balthica from the Gulf of Gdansk, neoplasia is considered as a serious epizootic linked to bad environmental conditions and high levels of pollutants. Previous research was focused on the diagnosis, prevalence, seasonality and histopathological characteristic of the cancer. This study is focused on electron microscopy analyses describing cell ultrastructure abnormalities related to neoplasia. Examinations using the electron microscopy highlighted changes confined to anatomic ultrastructures, shapes and functions of neoplastic cells. The lobulated appearance of the nucleus, changes in cellular matrix and the occurrence of large granular cells with hyperchromatic nuclei, atypical Golgi structures and deterioration of rough endoplasmic reticulum manifested the disease. The presence of atypical mitochondria, free ribosomes and hypertrophic nuclei suggests the adaptation of neoplastic cells to increased mitotic activity, while the observed modification of cellular membranes may reflect functional changes connected to increased pinocytotic activity or intercellular transport. The cancer cells were found to appear in two types, abnormal round-shaped cells and spindle-shaped cells, both with increased frequencies of cell division. Round-shaped cells typical for disseminated neoplasia were observed in all affected bivalves, in a few cases co-occurring with abnormal spherical cells. Spindle-shaped cells containing some intracytoplasmic filaments, and with a tendency of the nuclei to be orientated as in a palisade were interpreted as myofibroblasts-like cells and were observed in five out of eleven clams diagnosed as neoplastic. This finding represents the first demonstration of phenotypic differences in the cell types co-occurring in animals diagnosed as neoplastic and by that suggests coexistence of two types of bivalve cancer, disseminated neoplasia and probable fibrosarcoma. [Copyright &y& Elsevier]

Abstract: The responses of the burrowing bivalves Macoma balthica and Cerastoderma edule to chemical cues emitted by feeding shore crabs Carcinus maenas were investigated. M. balthica held in the laboratory and exposed to chemical signals in effluent water discharging from tanks containing C. maenas fed 20 M. balthica day−1 reacted by increasing their burial depths from approximately 30 mm to depths of >60 mm, over a period of several days. When the signal was removed the bivalves gradually returned to their original depth over 5 days. C. edule similarly exposed to effluent from crabs feeding on conspecifics showed no response. In an attempt to identify the signal inducing this burrowing response, M. balthica were exposed to a variety of chemical signals. Crabs fed M. balthica elicited the strongest response, followed by crabs fed C. edule. There were also small responses to effluent from crabs fed on fish, crabs previously fed on M. balthica and to crab faeces, but no responses to starved crabs, crushed M. balthica, or controls. We conclude that increased burrowing depth of M. balthica is induced by some as yet unidentified chemical cue produced by feeding crabs and is strongest when the crabs were fed on M. balthica. Unexpectedly, neither the presence of crabs themselves, nor of damaged conspecifics, was effective in eliciting a burrowing response. The mortality rates of M. balthica and C. edule selected by crabs when burrowed at normal depths and after exposure to effluent from feeding crabs were different. Crabs selected 1.5 times more C. edule than M. balthica when both species were burrowed at their normal depths, but 15 times more after the tanks had been exposed to effluent from feeding crabs for 5 days. The burrowing response of M. balthica thus appears to reduce mortality significantly by displacing predation pressure on to the more accessible C. edule. [Copyright &y& Elsevier]

Abstract: Metallothionein level (MT), and acetylcholinesterase (AChE), catalase (CAT) and glutathione-S-transferase (GST) enzyme activities in the bivalves Mytilus edulis and Macoma balthica were investigated for seasonal variations from an inshore and an offshore site in the northern Baltic Sea. All the biomarkers showed variability, following mostly a similar pattern at both sites. Relationships between biomarkers and environmental factors and protein concentration and weight of target tissues were examined. In M. edulis, GST activity was related to Secchi depth, while in M. balthica a correlation with near-bottom oxygen saturation was observed. AChE activity correlated with the weight of the foot tissue of M. balthica. In both species, an integrated biomarker index indicated a stressed condition during the spring/early summer period. Strong seasonal variability in temperature and a concentrated period of food availability in spring–both governing the reproductive cycle of the bivalves–probably explains most of the observed natural variability in biomarkers in this sea area. [Copyright &y& Elsevier]

The exploited feeding areas of Macoma baithica and Marenzeiieria viridis were experimentally quantified in laboratory conditions. Our feeding trials showed that M. baithica was able to feed at a much wider surface area than M. viridis. Moreover, in the presence of M. i'iridis the feeding area of M. baithica and its cumulative increase in time were significantly higher than with- out the polychaete. These results suggest that M. baithica is superior to M. viridis in terms of feeding. The experiment supports the earlier findings that the presence of M. baithica appears to be a key factor limiting the further expansion of M. viridis in the northern Baltic Sea. [ABSTRACT FROM AUTHOR]

Recent studies of the Baltic clam Macoma balthica (L.) from the southern Baltic (the Gulf of Gdansk) have revealed striking morphological, histological and cytogenetic features. Strong deformation of the shell, including elongation of the posterior end and the appearance of an easily visible flexure in this part, has been recorded. The population contribution of the deformed blunt shelled (“irregular”) clams ranged from 0% to 65% and tended to increase with depth. The morphologically “irregular” clams had higher accumulated tissue concentrations of trace metals (As, Ag, Cd, Pb, Cu and Zn), indicating a different metal handling ability. Adverse conditions in deeper water regions of the Gulf (e.g. hypoxia, hydrogen sulphide, elevated bioavailability of contaminants) have been suggested as inducers of the phenotypical changes (morphological deformation) in part of the population and, in parallel, of the specific physiological adaptations that result in higher metal accumulation in the “irregular” clams.Cytogenetic and histological analyses showed the presence of tumours in gill cells and digestive system of the affected clams, the prevalence of disseminated neoplasia ranging from 0% to 94% depending on the site. The disease was manifested by a modified karyotype (i.e. an abnormal number and morphology of chromosomes), a higher activity of nucleolar organizer regions (AgNORs), and tissue lesions (enlarged cells, actively proliferative with pleomorphic nuclei). Bottom sediments showed acute toxicity and have been proposed as a source of an initialising carcinogenic factor. However, none of the ecotoxicological studies provided was successful in the clear demonstration of a single (or multifactorial) agent that can account for the disseminated neoplasia. [Copyright &y& Elsevier]

1. Design considerations dictate that systems work most efficiently when the separate components have processing capacities adjusted to each other, a principle known as symmorphosis. Adjustments in response to a variable environment are most likely to occur at the interlace between the environment and integrated systems In deposit-feeding bivalves, the interlace of the alimentary canal with the foraging environment consists of a siphon (intake), gills (flow generation) and palps (particle sorting). 2. Among European populations of the Baltic Tellin, Macoma baithica, the gill-to-palp mass-ratio was positively correlated with the foraging environment, characterized by median grain size of the ambient sediment (R = 0·71). 3. In laboratory-reared M. balthica gill-to-palp mass-ratio responded plastically to changes in ambient sediment characteristics, resulting in large ratios in coarse sediment and small ratios in fine sediment. 4. In a common garden experiment, relative differences in gill-to-palp mass-ratios of laboratory-reared M. balthica from tour populations appeared similar to those in the field, indicating a hieritable component to the variation in integrated organ systems. 5. Phenotypically plastic adjustments ensure a close match between the integrated system of the alimentary canal to the local foraging environment. Heritable variation among populations may reflect different selective optima related to consistent differences in the environment between the populations. [ABSTRACT FROM AUTHOR]

In many estuaries worldwide, climate trends together with human diversion of fresh water have dramatically impacted the benthos. Such impacts have sometimes been complicated by exotic species, whose invasion and persistence can be mediated by wide variations in freshwater inflow. Monitoring such changes usually involves periodic samples at a few sites; but sampling that does not recognize variation at a range of spatial and seasonal scales may not reveal important benthic trends. San Pablo Bay, in northern San Francisco Bay, has extreme fluctuations in freshwater inflow. This bay also experienced a major benthic change with introduction of the Asian clam (Potamocorbula amurensis) in 1986. This species initially displaced the former community, but later appeared to vary in abundance depending on site and freshwater inflow. To investigate such patterns and provide guidelines for research and monitoring, we took 1746 core samples at six sites around San Pablo Bay from 19 October to 17 December 1999 and from 6 March to 19 April 2000. Most biomass consisted of the clams P. amurensis, Macoma balthica and Mya arenaria. Potamocorbula amurensis dominated the benthos at most sites in the fall and recruited a new cohort during winter, while there was weak recruitment in M. balthica and none in M. arenaria. At most but not all sites, densities of P. amurensis and M. arenaria declined dramatically over winter while M. balthica declined only slightly. The dominant clams had patch diameters >5 m at most but not all sites, and some showed inconsistent patch structure at scales of 100–1400 m. In this semiarid estuary with highly variable freshwater inflow, samples for research and monitoring should include multiple sites and seasons, and samples within sites should be ≥5 m apart to account for between-patch variation. Species abundance in winter 1999–2000 appeared to be affected by high freshwater inflows in 1997–1999, while spatial patterns were probably most affected by post-settlement dispersal and mortality. [Copyright &y& Elsevier]

The pre-spawning condition of adult bivalves is influenced by quantity and quality of available food. For bivalves, the essential polyunsaturated fatty acids (PUFAs) eicosapentaenoic acid (EPA) 20:5(n-3) and docosahexaenoic acid (DHA) 22:6(n-3) are presumed to determine the nutritional value of algae. Macoma balthica kept on a broodstock diet supplemented with PUFAs spawned a larger number of eggs (average 22220) per female and larger sized eggs (106.7 μm) compared to adults kept on a diet without PUFA supplementation (962.5 eggs with average size 99.8 μm).Larvae of M. balthica from the same parental pool however did not profit from a diet where a part was replaced with PUFA spheres. Instead, larvae reared on Isochrysis sp. showed lower mortality and higher growth rates than larvae fed on the same algae supplemented with lipid spheres. Crassostrea gigas larvae showed no clear response to a PUFA supplemented diet. [Copyright &y& Elsevier]

Suction dredging for cockles removes large cockles from tidal flats and may also cause mortality of non-target fauna and make the habitat less suitable for some species. This study examines whether suction dredging for cockles on tidal flats of the Dutch Wadden Sea had affected densities of non-target fauna, directly after fishing and one year later. Densities of non-target fauna in two randomly chosen undredged locations were compared to densities at the surrounding heavily commercially dredged area. A significant negative effect of cockle dredging on densities of 0-group Macoma balthica was observed and this effect persisted one year after dredging. The dredged area appeared to be less suitable for settlement of mussels Mytilus edulis. No significant effects of dredging on the mudsnail Hydrobia ulvae and on 0 and 1-group C. edule were found. For the mobile young Macoma balthica it seems unlikely that the effect found after one year was still due to the mortality caused by dredging and this suggests that the habitat was less suitable as a consequence of dredging. Thus, even in the highly dynamic ecosystem of the Wadden Sea, effects of bottom disturbance by cockle dredging may persist after one year. [Copyright &y& Elsevier]

In this laboratory study the effects of the cyanobacterium Nodularia spumigena (strain AV1) that produces hepatotoxic nodularin (NODLN), non-toxic Nodularia sphaerocarpa (strain UP16f) and purified NODLN on the infaunal clam Macoma balthica from the Baltic Sea were examined. N. sphaerocarpa (2.4 and 12.5 mg dw l−1), N. spumigena (4 and 20 mg dw l−1, intracellular NODLN content ca. 4 and 20 μg l−1) and purified NODLN (10 and 50 μg l−1) were applied in experimental tanks at 24 h intervals for 96 h. Water samples were taken during the experiment for the measurement of soluble NODLN concentrations. The concentrations of total hepatotoxins in the soft tissues were analysed with enzyme-linked immunosorbent assay (ELISA) and NODLN with high-performance liquid chromatography/diode array detection (HPLC/DAD). Acetylcholinesterase (AChE) activity was measured from the foot tissue samples taken at 0, 24 and 96 h. In the water phase, 60–100% of NODLN in the pure-toxin treatments and all the NODLN in N. spumigena treatments appeared as a yet unidentified form with NODLN-like spectral characteristics. The compound was present in similar quantities also in the non-toxic N. sphaerocarpa treatments. In the toxic N. spumigena treatments the tissue concentration of hepatotoxic NODLN-like compounds (measured with ELISA) increased from the control levels of 0.16 to 16.6 μg g−1 dw (24 h), reaching 30.3 μg g−1 dw at 96 h. However, <5% of the toxin detected by ELISA could be shown to be NODLN in HPLC/DAD analysis. Matrix-assisted laser desorption/ionisation-time of flight mass spectrometry (MALDI-TOF/MS) analyses revealed no NODLN-glutathione (GSH) conjugates in the tissues of M. balthica. Combining the responses in the AChE activity recorded after 24 and 96 h exposure, increases by 46% (N. spumigena) and 36% (soluble NODLN) compared with the control activity were observed in the low-level toxic exposures. Oppositely, decreases by 19% (N. spumigena) and 27% (soluble NODLN) of control activity were observed in the high-level exposures. Only the differences between the increased and decreased AChE activities were statistically significant, with individuals exposed to high levels expressing approximately 55% of the activity of those exposed to low concentrations. The results show that M. balthica readily ingests toxic N. spumigena and that accumulation of peptides takes place rapidly, which has potential food chain effects through toxin enrichment. However, it appears that M. balthica is at least partly able to metabolise NODLN. In addition to hepatotoxicity, NODLN seems to induce concentration-dependent neurotoxic effects; this must be taken into consideration when applying AChE activity as a biomarker of specific anthropogenic contamination (e.g. organophosphate and carbamate pesticides). [Copyright &y& Elsevier]

Abstract Mitochondrial DNA sequence data for 295 individuals of the marine bivalve Macoma balthica (L.) were collected from 10 sites across the European distribution, and from Alaska. The data were used to infer population subdivision history and estimate current levels of gene flow. Inferred historical biogeography was expected to be congruent with colonization of the Atlantic Ocean from the Pacific Ocean after the opening of the Bering Strait 3.5 Ma. In addition, the last glacial maximum, about 18 000 years ago, was expected to have been responsible for most of the present-day distribution of molecular variation within Europe, because the area must have been recolonized after confinement to France and the south of the British Isles during the last glacial maximum. Current gene flow was hypothesized to be high, because the larvae of M. balthica spend 2–5 weeks drifting in the water column. The geographical distribution of one highly diverged haplotype clade was found to be disjunct and was encountered exclusively in samples from the Baltic Sea and Alaska. A molecular clock calibration for marine bivalve cytochrome-c -oxidase I dates this clade as having split off from the other haplotypes 9.8–39 Ma. Multiple colonizations of the Atlantic Ocean from the Pacific by M. balthica may explain the strong differences found between Baltic Sea and other European populations of this species. The sympatric occurrence of the highly diverged mitochondrial lineages in western parts of the Baltic Sea points to secondary admixture. With the use of coalescent analysis, population divergence times for French vs. other non-Baltic European populations (‘Atlantic population assemblage’) were estimated at a minimum of about 110 000 years ago, well before the last glacial maximum 18 000 years ago. Signatures of population divergence of M. balthica that appear to have originated during the Pleistocene have thus survived the last glacial maximum. Some of the... [ABSTRACT FROM AUTHOR]

Many soft-bottom benthic invertebrates display sediment-associated patterns of dispersion. Habitat selection experiments have shown that the larvae of some species can choose to settle in favourable habitats, and this process could establish patterns observed in the field. However, many soft-bottom infauna, including the bivalves Cerastoderma edule and Macoma balthica, show postlarval relocation. Such movements could obviate patterns established at initial settlement, and active habitat selection in these postlarval stages may be responsible for adult distributions. We investigated sediment selection in postlarval stages of these bivalves on an intertidal mudflat using a field survey and experiment. Our 1225 m2 grid survey showed a strong spatial correlation between the smallest individuals of both species, which showed no correlation with sediment characteristics. Larger size groups showed increasingly strong associations with sediment type: C. edule with sandy sediments and M. balthica with muddy. In our sediment choice experiment, the numbers of bivalves recruited into muddy, sandy and muddy depression (to detect small-scale hydrodynamic effects) treatments after 2 days in the field were recorded. There were no significant treatment effects in either C. edule or M. balthica. All bivalves were too large to be initial settlers; they were thus relocating individuals. Neither the survey nor the experiment provided any evidence that relocating individuals of these two species could choose the sediment in which they alight. The sediment-related patterns observed in adults of these species must therefore be due to other mechanisms, possibly postsettlement predation by Carcinus maenas and/or an ability to leave less-preferred sediments in a series of migrations until the preferred habitat is found. [Copyright &y& Elsevier]

Fourteen acidic and neutral free amino acids (FAA) were investigated in soft tissue of Macoma balthica from different depth zones of the Gulf of Gdansk (Baltic Sea) over a full seasonal cycle. The dry weight of the bivalves and physico-chemical parameters of overlying bottom water and surface sediments were measured simultaneously at each site. In the brackish waters of the Baltic, the main pool of FAA is composed of Ala, Gln, Arg, Gly and Orn which represent approximately 80% of the total. Compared to the full saline environments, the composition of FAA in the clams from the Baltic differs substantially. The differences can be attributed to the lower salinity of the Baltic. In the Baltic, Gly appears to play a most important role in regulating intracellular osmolarity in the clams, a function performed primarily by Tau in Atlantic and North Sea populations. Spatio-temporal variations of the FAA are affected by biotic and environmental parameters; their respective influence differs with the amino acids. The concentration of Arg depends on its uptake from the external medium. However, its level might be temporarily modified by stress-induced metabolic transformation (e.g. hydrolysis to Orn) caused by changes in the ambient environment. The concentration of Ala increases with depth, probably because of physiological adaptations of the animal to diminishing oxygen concentration through anaerobic glucose catabolism. Biosynthesis of Ala, similarly to Gln, in the shallower zone is generally related to the physiological state of an organism. The concentration of Gly is most likely regulated by internal mechanisms driven by gonadal development and reproduction. [Copyright &y& Elsevier]

Abstract Understanding the origin, maintenance and significance of phenotypic variation is one of the central issues in evolutionary biology. An ongoing discussion focuses on the relative roles of isolation and selection as being at the heart of genetically based spatial variation. We address this issue in a representative of a taxon group in which isolation is unlikely: a marine broadcast spawning invertebrate. During the free-swimming larval phase, dispersal is potentially very large. For such taxa, small-scale population genetic structuring in neutral molecular markers tends to be limited, conform expectations. Small-scale differentiation of selective traits is expected to be hindered by the putatively high gene flow. We determined the geographical distribution of molecular markers and of variation in a shell shape measure, globosity, for the bivalve Macoma balthica (L.) in the western Dutch Wadden Sea and adjacent North Sea in three subsequent years, and found that shells of this clam are more globose in the Wadden Sea. By rearing clams in a common garden in the laboratory starting from the gamete phase, we show that the ecotypes are genetically different; heritability is estimated at 23%. The proportion of total genetic variation that is between sites is much larger for the morphological additive genetic variation ( Q ST = 0.416) than for allozyme ( F ST = 0.000–0.022) and mitochondrial DNA cytochrome- c -oxidase-1 sequence variation (Φ ST = 0.017). Divergent selection must be involved and intraspecific spatial genetic differentiation in marine broadcast spawners is apparently not constrained by low levels of isolation. [ABSTRACT FROM AUTHOR]

Higher than average recruitment among bivalves on the intertidal flats of the Wadden Sea was often observed after severe winters in the period 1940–1995. The occurrence of another severe winter in 1995/96 prompted us to test the hypothesis of severe winters leading to universally high bivalve recruitment on a large geographic scale (500 km coastline) in temperate shallow waters. We analysed data sets on bivalve abundance from seven areas in the Dutch, German and Danish Wadden Sea. The longer data sets showed generally higher bivalve recruitment in the 1970s and 1980s than in the 1990s which may be related to the near absence of severe winters since 1987. Considering the period 1988 onwards (the longest possible period in which 1995/96 was the only severe winter), recruitment of Cerastoderma edule was in 1996 - in agreement with our hypothesis - above average in all seven areas investigated. In contrast, recruitment of Macoma balthica and Mya arenaria was for the same period above average only in the southern Wadden Sea (south-west of Jade Bay) but not in the northern Wadden Sea (north of Eiderstedt peninsula). These regional differences may be related to (i) the different topography of the northern Wadden Sea (with barrier islands west of the mainland) compared to the southern Wadden Sea (with barrier islands north of the mainland) and subsequent differential effects of wind-induced currents on bivalve recruitment, (ii) differences in biotic factors such as standing stocks, larval supply or epibenthic predation or (iii) changes in environmental conditions. Our results demonstrate that large-scale comparisons along coasts are an indispensable addition to insights derived from local studies alone. [Copyright &y& Elsevier]

The parasite manipulation hypothesis predicts that parasites should be selected to manipulate host behaviour to facilitate transmission to the next host. The bivalve Macoma balthica burrows less deep when parasitized by the trematode Parvatrema affinis. Shallow burrowing increases the likelihood of ingestion by birds, their final hosts, and therefore this has been interpreted as manipulation by the parasite. When unparasitized, M. balthica displays seasonal changes in burrowing depth, becoming less accessible to predators in winter. If shallow burrowing of parasitized individuals is due to direct manipulation by the parasite, the availability of parasitized individuals should be high throughout the year, or at least especially in the season when most birds are present and potential transmission rates are highest. We compared burrowing depths of parasitized and unparasitized individuals in a single population during seven consecutive years. Parasitized individuals showed reduced burrowing depths but, in contrast to the prediction, the effect of parasites on availability to predators was smallest, not largest, in the season with the highest bird numbers. The parasite P. affinis competes for energy with the host, and M. balthica with low energy stores are known to reduce depth of burrowing. When we included size-corrected somatic ash-free dry mass (as an estimate of the energy stores) in our statistical analysis, the effect of infection on burrowing depth disappeared. Thus the effect of infection on burrowing depth is likely to be an unavoidable, indirect effect of the channelling of energy towards the parasite, causing the starving individual to take greater risks in the acquisition of food. Since both the seasonal pattern and the magnitude of increased availability of parasitized individuals are inadequate, the increased exposure of parasitized M. balthica to the final host does not seem to represent an example of adaptive host manipulation by the parasite. [ABSTRACT FROM AUTHOR]

The bivalve Macoma balthica is a common species in the Wadden Sea and North Sea. Juveniles temporarily use nurseries in the high intertidal. To explain this nursery use, predation pressure was examined for both juvenile and adult Macoma at low and high tidal flats. The study was carried out in the eastern Dutch Wadden Sea.Shrimps Crangon crangon, adult crabs Carcinus maenas, gobies Pomatoschistus and juvenile flatfish were more abundant and larger on low than on high tidal flats, but 0-group Carcinus was more abundant on the high tidal flats. Crangon and 0-group Carcinus stomachs frequently contained Macoma remains. These predators selectively preyed on small 0-group Macoma, both in the field and in laboratory experiments. The effect of predation by epibenthic animals and birds, on the low and high tidal flats, was examined in exclosure experiments (2 mm mesh). There was no effect of epibenthos exclosure on adult Macoma. For 0-group Macoma, densities were higher in exclosures than in the controls where predators had normal access. The density reduction by epibenthic predators was much larger in the low than in the high intertidal. We found no effect of bird predation on densities of 0- and 1+group Macoma.Thus, 0-group Macoma is under high predation pressure by epibenthos in the low intertidal, especially by shrimps, while they are relatively safe in the high intertidal. However, most of the shellfish outgrow their epibenthic predators during their first summer. Therefore, it becomes safe for the bivalves to redistribute to locations where epibenthic predators are abundant, during their first winter. On the other hand, it did not become clear from this study why many of the larger Macoma leave the high intertidal. Concluding, the nursery use of Macoma-spat in the high intertidal is probably, at least partly, an adaptation to avoid epibenthic predation. [Copyright &y& Elsevier]

As a characteristic buried tellinid bivalve, Macoma balthica has a long inhalent siphon that enables it to feed in two different ways: deposit and suspension feeding. To deposit feed efficiently on benthic microalgae, Macoma has to live close to the sediment surface, where it can graze an extensive surface area, but is within reach of many predators. Individuals that are more safely buried at a greater depth can only suspension feed, or deposit feed from a small surface area. We expected local differences in burying depth on intertidal mudflats to be caused by differences in immersion time (i.e. time available for feeding, particularly suspension feeding), since immersion time has been shown experimentally to affect body condition positively, and since body condition and burying depth in Macoma are postively related in the field. To test this we experimentally manipulated immersion time, and followed changes in burying depth and body condition. In the experiments, longer immersion time went consistently with greater burying depth of Macoma and higher body condition. On a transect in the western Wadden Sea, the deepest Macoma were indeed found at the intertidal level with the longest immersion time, but these were at that time not the animals with the highest body condition. Within each locality, however, body condition was positively correlated with burying depth. The experimental data and the within-locality data support the hypothesis that longer immersion time may influence burying depth through body condition. However, the fact that between-locality differences in burying depth seemed to be consistently related to immersion time, but not to body condition, indicates that body condition alone does not explain place-to-place variation in burying depth. [Copyright &y& Elsevier]

Juveniles of the bivalve Macoma balthica live on tidal flats in the Wadden Sea. This study examined the interaction of Macoma with the infaunal polychaetes Arenicola marina and Nereis diversicolor and the gastropod Retusa obtusa. The distribution of M. balthica spat on the flats, shortly after settlement in April, showed a positive correlation with the Arenicola distribution and a negative correlation with Nereis distribution. There were no locations where Macoma spat and Retusa occurred together. In August, Macoma spat had grown too large for predation by intertidal infauna.Small individuals of Macoma spat were found in stomachs of Arenicola (0.14 worm−1) and Nereis (0.05 worm−1). Laboratory experiments showed that Nereis and Retusa could reduce Macoma spat abundance, both in the absence and presence of sediment and alternative prey. Arenicola reduced the abundance of small Macoma (<1 mm) in sediment without, but not with, alternative prey.In field experiments, we manipulated the density of Arenicola in 0.25–1 m2 plots and of Nereis in 0.03 m2 cages and examined the effect on Macoma density several weeks later. We found a significant negative relation between densities of polychaetes and Macoma spat for both polychaete species in these experimental plots. Peculiarly, we found a significant positive relation between manipulated Nereis density and adult Macoma density in the cages; we cannot explain this.Consumption rates, calculated both from stomach contents and from field experiments, were 45 to 102 Macoma m−2 d−1 for Arenicola and 5 to 116 Macoma m−2 d−1 for Nereis. These values are higher than recorded consumption rates by epibenthic predators in the same area. Nevertheless, between-year differences in year-class strength could not be explained by differential abundance of these polychaetes.In conclusion, Arenicola and Nereis had a negative effect on the abundance of Macoma <1.5 mm, which was at least partly caused by direct consumption. Retusa obtusa can eat juvenile Macoma, but probably did not so in the study area, because there were no locations where Retusa and Macoma spat occurred together in the period that Macoma was <2 mm. [Copyright &y& Elsevier]

The Copper River Delta, Alaska is the largest contiguous coastal wetland system along the West Coast of North America. Vast expanses of tidal mud flats formed by sediments carried by the suspended load of the Copper River serve as a connection between the Gulf of Alaska and the extensive network of wetlands, rivers and sloughs of the delta system. In addition to providing habitat for resident fish, shrimp and crabs, these tidal flats serve as critical feeding grounds for up to 5 million migratory shorebirds as well as an entry and exit corridor for three species of commercially fished salmonids. Here we report the first description of the benthic community of these intertidal flats. Between April and September 2000, we conducted three samplings on the Copper River Delta in which we quantified benthic macro-invertebrates inhabiting silt-clay sediments, the dominant substrate in the system, over a range of tidal inundation. Specifically, sampling was performed in two areas on the delta: near the outflows of the Eyak River and Pete Dahl Slough. Pore-water salinity of surficial sediment ranged from 4 psu during peak summer flow of the Copper River to 14 psu in April prior to increased riverine input. Sediment temperatures corresponded to ambient air temperatures with lowest temperatures during the April–September observation period recorded in April (4°C) and warmest in August (16°C). The benthic community of the delta''s tidal flats was characterised by low species diversity and was dominated by the tellinid bivalve Macoma balthica, which reached densities greater than 4000 m−2. Age–length relationship of M. balthica indicated slow growth and longevity of up to 8 years. Polychaete densities, primarily the phyllodocid Eteone longa, were low throughout the study period, reaching a maximum of only 700 m−2 in August. Amphipod densities, primarily the corophid amphipod Corophium salmonis, were high (up to 7000 m−2) only during the August sampling. Spatial patterns of benthic invertebrate abundance were best explained by differences in tidal inundation with longer inundation corresponding to greater invertebrate densities. Temporal changes in abundance of polychaetes, amphipods, and M. balthica recruits corresponded to increases in sediment temperatures. Natural or human-induced changes to M. balthica populations could impact the food web of the delta, which could cascade to larger geographic impacts because of the importance of the delta to migratory species. [Copyright &y& Elsevier]

At large spatial scales (>1 km), the forces structuring marine communities are diverse and include oceanographic, geological, and human processes, as well as availability of regional species pools. Few studies in marine and estuarine systems have examined the joint effects of predation (top-down) and food availability (bottom-up) in controlling populations at these scales. We compared the relative influence of top-down and bottom-up factors to population abundance of the infaunal clam Macoma balthica in two ecosystems differing in spatial extent by an order of magnitude: the York River, 50 km long, and the Rhode River, 5 km long. In both rivers, two habitat types could be differentiated by either high or low density of Macoma. In the York, Macoma abundance was tenfold greater upriver than downriver, predator abundance was greater in the high clam density habitat (i.e. upriver), and predation was only evident where clams were dense (i.e. upriver, where food availability for predatory crabs was elevated). In this large-scale system, predators were apparently unable to travel between upriver and downriver habitats where food availability differed. Moreover, sedimentary carbon (i.e. food for deposit-feeding clams) was significantly greater in the high clam-density habitat; this is consistent with the hypothesis that bottom-up factors principally dictate clam density, and moreover, that clam density drives predator density. In the Rhode River, a smaller-scale system, predators could easily move among habitats differing in clam density. Macoma abundance and predator abundance were both threefold greater in mud than sand. In contrast, predation intensity was higher in sand than mud, consistent with a hypothesis of top-down control. As in the large-scale system, sedimentary carbon was significantly greater in the high clam-density habitat (i.e. mud). In this smaller scale system, both top-down and bottom-up factors contributed to habitat-specific patterns in clam population abundance. Hence, in this marine benthic ecosystem, bottom-up control was important at both large and small spatial scales, whereas top-down control was only important at the smaller spatial scale. [ABSTRACT FROM PUBLISHER]

Abstract 1. The importance of recruitment processes in determining benthic population dynamics has received considerable interest among marine ecologists in the last two decades. Observational demographic studies, in which recruits were followed to and through the adult state, aimed to estimate whether the variation in the numbers of recruits is dampened by density-dependent post-recruitment processes. These studies revealed conflicting results on the importance of post-recruitment density dependence, but were performed over at most a few years. 2. Based on a study of the demography and growth of the bivalve Macoma balthica (L.) on an intertidal flat in the Dutch Wadden Sea for a period of almost 30 years, we explored the extent to which the among-cohort variability in recruitment was reflected in the secondary production (which results from the combined action of recruitment, mortality and growth) of the adult population. 3. Since growth in length was of the Von Bertalanffy type and post-recruitment instantaneous mortality rate was approximately constant for each cohort, production could be explicitly written in terms of the following parameters: recruit density, condition (ratio of mass to cubic length), Von Bertalanffy growth coefficient, asymptotic cubic length, and expected life span. 4. Decomposing the among-cohort variability in production in terms of the covariance matrix of these parameters revealed that most of the variance in production could be attributed to year-to-year variability in recruit density. 5. Small negative covariances between recruit density and asymptotic size and expected life span indicated weak density dependence after recruitment. 6. The major cause of temporal variation in abundance and production is inter-annual variation in recruitment and competition for resources seems to play a small role in affecting post-recruitment population density and production. [ABSTRACT FROM AUTHOR]