Herman Hummel

Geographic and seasonal patterns and limits on the adaptive response to temperature of European Mytilus spp. and Macoma balthica populations

Seasonal variations in seawater temperature require extensive metabolic acclimatization in cold-blooded organisms inhabiting the coastal waters of Europe. Given the energetic costs of acclimatization, differences in adaptive capacity to climatic conditions are to be expected among distinct populations of species that are distributed over a wide geographic range. We studied seasonal variations in the metabolic adjustments of two very common bivalve taxa at European scale. To this end we sampled 16 populations of Mytilus spp. and 10 Macoma balthica populations distributed from 39° to 69°N. The results from this large-scale comprehensive comparison demonstrated seasonal cycles in metabolic rates which were maximized during winter and springtime, and often reduced in the summer and autumn. Studying the sensitivity of metabolic rates to thermal variations, we found that a broad range of Q10 values occurred under relatively cold conditions. As habitat temperatures increased the range of Q10 narrowed, reaching a bottleneck in southern marginal populations during summer. For Mytilus spp., genetic-group-specific clines and limits on Q10 values were observed at temperatures corresponding to the maximum climatic conditions these geographic populations presently experience. Such specific limitations indicate differential thermal adaptation among these divergent groups. They may explain currently observed migrations in mussel distributions and invasions. Our results provide a practical framework for the thermal ecophysiology of bivalves, the assessment of environmental changes due to climate change and its impact on (and consequences for) aquaculture.

Response of the blue mussel Mytilus edulis L. following exposure to PAHs or contaminated sediment

Abstract The polycyclic aromatic hydrocarbons fluoranthene and benzo(a)pyrene significantly reduced the feeding rate of mussels. For both compounds the tissue concentration resulting in a 50% reduction of the clearance rate (TEC50) was calculated. At high tissue concentrations both aromatic compounds reduced the tolerance of mussels to aerial exposure, whereas at low tissue concentrations an improved response was noticed. The activities of the antioxidant enzymes Superoxide dismutase (SOD) and catalase were elevated only at low tissue concentrations of fluoranthene and benzo(a)pyrene. At the highest measured tissue concentrations the activity of both enzymes was reduced, possibly due to a narcotic effect. The reproductive success rate of mussels appeared to be affected negatively by the investigated hydrocarbons. The results of a pilot experiment indicate that mussels can be used also for the testing of contaminated sediments.

“Survival in air” of the blue mussel Mytilus edulis L. as a sensitive response to pollution-induced environmental stress

Mussels, Mytilus edulis, were exposed for periods of 6 weeks at various locations in Dutch coastal waters during 1989 and 1990. “Survival in air” showed to be a sensitive response parameter for indicating pollution induced environmental stress in transplanted mussels sampled from eight field sites. Increased tissue contaminant levels, especially PCBs and PAHs, correlated with a reduced survival time during aerial exposure. Three weeks exposure of mussels in the laboratory to 1 μg · 1−1 PCBs affected the aerial survival time negatively. Laboratory experiments did not indicate that lowered salinity influences the “Survival in air” response after sufficient acclimation (15 days), facilitating the use of this response parameter in both marine and estuarine waters.

Spatial and seasonal differences in the PCB content of the mussel Mytilus edulis

Seasonal and spatial variation in the concentration of PCBs, fats (non-polar lipids) and total lipids and the condition of the mussel Mytilus edulis were assessed in three differing water bodies of the Dutch delta area. Highest concentrations of PCBs in the mussel were found in the Westerschelde estuary, with much lower concentrations in the Oosterschelde and the brackish lake Grevelingenmeer. Spatial differences were strongly related to salinity; lower concentrations were found at the more saline stations, pointing to freshwater inputs as being the origin of the PCBs in mussels. The PCB concentration in mussels, on the basis of dry or total weight, in general increased during summer, autumn and winter and decreased strongly during spring. The strong decrease is related to the spawning of gametes. Seasonal changes in the PCB concentration on the basis of the dry weight were not related to changes in the fat content or the condition of the animals. Only PCBs on a fat basis were negatively related to fat content, indicating a dilution of PCBs during seasonal fat accumulation and concentration of PCBs during fat utilization, in such a way that the total PCB concentration in the animal remains the same. It seems that, besides reproduction, equilibrium partitioning is the most probable mechanism that determines the PCB content of mussels.

Uniform variation in genetic traits of a marine bivalve related to starvation, pollution and geographic clines

Consistent patterns of genetic variation in the marine bivalve Macoma balthica (L.) were found after exposure to low levels of copper, starvation, and along geographic clines. The geographic clines were related to temperature and salinity. Genetic differences were primarily found in the LAP (Leucine aminopeptidase) locus; under stress the frequency of one specific allele, and thereby the heterozygosity, decreased strongly. The degree of changes depended on age and condition of the animals; adult animals showing stronger changes than juveniles. Low concentrations of copper yielded stronger genetic changes than high concentrations. It is concluded that genetic diversity and stress-sensitivity of species change predictably along a gradient in response to environmental harshness. Animals near the southern limit of their range were more sensitive to additional stress.

The glycogen content in stressed marine bivalves: The initial absence of a decrease

Abstract 1. 1. Changes in the glycogen content, condition, stomach content and acetic acid concentration of mussels Mytilus edulis and cockles Cerastoderma edule were followed during periods of up to 14 days of exposure (to air) at temperatures of 5 and 20°C. 2. 2. In animals with a high glycogen content the glycogen is not used during the first 3 to 7 days, at high and low temperature respectively. 3. 3. After this latent period the glycogen concentration often decreased, coinciding with a high mortality and an increase of the concentration of acetic acid. 4. 4. In cockles with a low glycogen content, and kept at a high temperature, glycogen can be used from the beginning of the stress period. 5. 5. Between species no clear differences were found. 6. 6. The stomach content decreased during exposure; however, the stomach content amounted to only 0.5 to 0.7% of the body weight, and is thought to be of minor importance as an energy source during the stress period. 7. 7. Especially at the higher temperatures glycogen finally is transformed into acetic acid. 8. 8. It is concluded that during exposure, the animals do not die because of a lack of energy reserves, but because of a high accumulation of acids.

Distribution of dissolved and labile particulate trace metals in the overlying bottom water in the Vistula River Plume (Southern Baltic Sea)

Overlying bottom water samples were collected in the Vistula River plume, southern Baltic Sea, (Poland) and analysed for dissolved and labile particulate (1 M HCl extractable) Cu, Pb, Zn, Mn, Fe and Ni, hydrological parameters being measured simultaneously. Particulate organic matter (POM), chlorophyll a and dissolved oxygen are key factors governing the chemical behaviour of the measured metal fractions. For the dissolved Cu, Pb, Zn, Fe and Ni two maxima, in the shallow and in the deeper part of the river plume, were found. In the shallow zone desorption from seaward fluxing metal-rich riverine particles account for markedly increased metal concentrations, as confirmed also by high particulate metal contents. For Pb, atmospheric inputs were also considered to have contributed to the elevated concentrations of dissolved Pb adjacent to the river mouth. In the deep zone desorption from detrital and/or resuspended particles by aerobic decomposition of organic material may be the main mechanism responsible for enrichment of particle-reactive metals (Cu, Pb, Zn) in the overyling bottom waters. The increased concentrations of dissolved Fe may have been due to reductive dissolution of Fe oxyhydroxides within the deep sediments by which dissolved Ni was released to the water. The distribution of Mn was related to dissolved oxygen concentrations, indicating that Mn is released to the water column under oxygen reduced conditions. However, Mn transfer to the dissolved phase from anoxic sediments in deeper part of the Vistula plume was hardly evidenced suggesting that benthic flux of Mn occurs under more severe reductive regime than is consistent with mobilization of Fe. Behaviour of Mn in a shallower part has been presumably affected by release from porewaters and by oxidization into less soluble species resulting in seasonal removal of this metal (e.g. in April) from the dissolved phase. The particulate fractions represented from about 6% (Ni) and 33% (Mn, Zn, Cu) to 80% (Fe) and 89% (Pb) of the total (labile particulate plus dissolved) concentrations. The affinity of the metals for particulate matter decreased in the following order: Pb > Fe > Zn > or = > Cu > Mn > Ni. Significant relationships between particulate Pb-Zn-Cu reflected the affinity of these metals for organic matter, and the significant relationship between Ni-Fe reflected the adsorption of Ni onto Fe-Mn oxyhydroxides. A comparison of metal concentrations with data from other similar areas revealed that the river plume is somewhat contaminated with Cu, Pb and Zn which is in agreement with previous findings on anthropogenic origin of these metals in the Polish zone of southern Baltic Sea.

A comparative study on the relation between copper and condition in marine bivalves and the relation with copper in the sediment

The relation between condition and copper concentration was assessed in three bivalve species (the Baltic clam Macoma balthica, the cockle Cerastoderma edule and the mussel Mytilus edulis) from several Dutch and French estuaries. In general, the copper concentration in the bivalves was negatively related to condition. The slope of this relation was strongest in the Baltic clam M. balthica, with an almost inversely proportional change of Cu concentration at a change of condition, indicating an almost constant Cu content (body burden) per individual, i.e. no elimination or accumulation of Cu. In the mussel M. edulis, the slope of this relation was absent or the weakest, with an almost uniform Cu concentration at a change of condition, indicating an almost proportional elimination (or accumulation) of Cu at a decrease (or increase) of the condition. Thus in Baltic clams the copper body burden is kept at an almost constant level, whereas in mussels the copper concentration is almost constant. In Baltic clams, but not in mussels and cockles, spatial differences were related to copper concentrations in the sediment. It is discussed that mussels maintain a homeostatic Cu concentration within a short time-period (days; copper highly exchangeable by complexation to metallothioneins), whereas the Cu concentration in Baltic clams is more slowly regulated (several weeks; fossilization of copper in mineral granules) and remains on an average in (partitioning) equilibrium with the Cu concentration in the sediment. Therefore, the Baltic clam is a better indicator of sediment copper pollution than the commonly used mussel. In addition, the observed differences between species were compared with differences in their life-strategies. The Cu concentration in the sediment is strongly related to the silt fraction (< 16 μm) of the sediment.

Genetic traits in the bivalve Mytilus from Europe, with an emphasis on Arctic populations

Abstract Genetic and some ecophysiological traits of mussels collected in the European Arctic, up to their northeastern distribution limit in the Barents Sea, were studied and compared with traits of mussels from the Mediterranean, Atlantic and Baltic. The genetic traits of these populations were analysed by isoenzyme electrophoresis on seven loci in order to assess the Mytilus complex to which populations in the Arctic region belong. Ecophysiological variables, the weight-index and glycogen were analysed to assess the physiological fitness of the populations. Three distinct groups were recognised: (1) Mytilus (edulis) galloprovincialis in the Mediterranean and Spain, (2) M. (edulis) edulis along the Atlantic coast from the Netherlands northwards into Russia, and (3) the Baltic Mytilus (edulis) trossulus. The mussels from populations in the Russian Arctic all belong to the Atlantic Mytilus (edulis) edulis group. The genetic variability and ecophysiological measures indicated that the sub-Arctic White Sea mussel populations have a relatively lower performance capacity, whereas those in the Arctic at the edge of their northern distribution showed a surprisingly strong performance.

The respiratory performance and survival of the bivalve Macoma balthica (L.) at the southern limit of its distribution area: a translocation experiment

The hypothesis was tested that animals near their extreme Southern limits, due to high temperatures, have a high respiration rate, whereby they reach an extreme low weight-index and ultimately disappear. At estuarine stations the respiration rate of Macoma balthica (L.) (Baltic clam) did not show interseasonal changes, indicating seasonal acclimation, but within the season the respiration increased with increasing temperature, indicating the absence of short-term acclimation. In clams translocated from the Netherlands towards the Bidasoa estuary, 200 km South of their Southern distribution limit, their respiration rate was higher and weight-index lower than in specimens living in Dutch estuaries. Irrespective of an effect of the temperature, clams exposed in experiments to water from Bidasoa showed a higher respiration than clams exposed to water from the other stations. Moreover, at non-estuarine stations with a low food content, the clams showed reversed acclimation, i.e., the respiration rates in winter were much lower than summer rates, most probably a strategy to conserve energy by means of a depressed metabolism. A weight index of 5 mg DW/cm(3) and glycogen content of 2% DW are suggested as the minimal values below which the metabolic energy balance of Baltic clams becomes more negative and the clam population disappears. It was concluded that factors other than temperature influenced the respiration and weight-index of clams, and hence their presence or absence, e.g., food concentration, innate seasonal cycles, and possible pollutants in the water.