Marenzelleria cf. viridis and the sulphide regime

To find out how the polychaete Marenzelleria cf. viridis could spread successfully into the habitat of the Darss-Zingst Bodden Chain, one important environmental factor for sediment dwelling animals was examined: hydrogen sulphide. To investigate the stress of this environmental factor, hydrogen sulphide was continuously examined in the pore water of the sediment and burrows of M. cf. viridis. Metabolic activity was recorded by direct and indirect calorimetry. Depending on water temperature, organic matter content of the sediment and salinity, the sulphide concentration in the pore water varied between 1.5 and 4.2 mmol l^-1 being high during summer and in winter when the sediment and overlying water was ice covered. In microcosm experiments water of M. cf. viridis-burrows showed variations in sulphide between 145 and 210 µmol l^-1 but pore water concentration was much higher (6.5 mmol l^-1). In the presence of oxygen animals exhibited an accelerated metabolic rate which was met by a fully aerobic metabolism at Po₂ of 20 to 7.5 kPa and sulphide concentration of 215–245 µmol l^-1. When oxygen is absent the heat production was only slightly elevated (103%) when compared to the anoxic control. The elevated heat production of the animals during sulphide exposure and oxygen may be due to detoxification processes. In this case thiosulphate is formed probably via mitochondrial oxidation and therefore may account for additional ATP-gain.     

Factors controlling the distribution of Marenzelleria cf. viridis, Pygospio elegans and Streblospio shrubsoli (Polychaeta: Spionidae) in the southern Baltic Sea, with special attention for the response to an event of hypoxia

The large-scale distribution patterns of the estuarine spionid polychaetes Marenzelleria cf. viridis, Pygospio elegans, and Streblospio shrubsoli were studied in the Pomeranian Bay (southern Baltic Sea) at 20 stations in April 1993 and their relationship with hydrographic factors and sediment composition investigated. The densities of M. cf. viridis and S. shrubsoli decreased rapidly with increasing distance offshore. A corresponding decrease in phytoplankton concentration offshore is suggested as the main cause for the observed spatial patterns. P. elegans was more evenly distributed. At 10 selected stations further samples were collected between April 1993 and July 1996 to study the response of abundance and biomass of the three polychaetes to changes in environmental conditions and to investigate interspecific interactions within the macrofauna. Salinity, pelagic food supply, and sediment parameters did not change. At three of these stations redox potential of the sediment and macrofauna composition were affected by an unusual hypoxic/anoxic event in July/August 1994. All three spionid species were able to survive moderate seasonal hypoxic conditions, but died after exposure to severe anoxia. Special attention was focused on the re-establishment of the spionids. The rate with which M. cf. viridis and P. elegans re-colonised defaunated stations varied between a few weeks and two years. These differences in rate of re-establishment were attributed to the distance from undisturbed recruitment areas, and to the severity of the oxygen deficiency. Increasing densities of P. elegans and S. shrubsoli after the hypoxic event coincided with a reduced abundance of the bivalve Mya arenaria, suggesting a negative interspecific interaction.     

Marenzelleria cf. viridis (Polychaeta: Spionidae) – ecophysiological adaptations to a life in the coastal waters of the Baltic Sea

Since its introduction into the Baltic Sea about ten years ago, the polychaete species Marenzelleria cf. viridis has spread rapidly. Meanwhile, this spionid settled in remarkable numbers predominantly in the coastal waters which provide some of the more variable and unstable habitats in terms of abiotic conditions. In the present paper, some of the underlying biochemical and physiological processes were experimentally examined which enables this polychaete to deal with several kinds of environmental stress such as variations in salinities, low oxygen concentrations and occurrence of hydrogen sulphide. The results obtained reveal that in the process of acclimation to different salinities free amino acids are involved, mainly alanine and glycine. The content of these amino acids were increased in worms acclimated to a higher salinity. When exposed to low ambient oxygen concentrations (severe hypoxia) M. cf. viridis started to produce energy anaerobically via pathways known from several other marine organisms. The presence of hydrogen sulphide in higher concentrations (1 mmol l^-1) results in a more rapid production of succinate (indicator for anaerobic energy production via the succinate-propionate pathway) as compared to hypoxia alone, indicating a higher energy demand. Nevertheless, when exposed to enhanced ambient hydrogen sulphide concentrations (3 mmol l^-1), M. cf. viridis is less affected by this toxic compound than other marine species, such as the related species Marenzelleria cf. wireni (North Sea). The field data indicate that the metabolic response obtained under experimental conditions is also important in the worm's natural habitat. Although the extent of anaerobic metabolites produced as well as the amount of thiosulphate (main sulphide detoxification product) was lower in the field as compared to the experiments, M. cf. viridis very likely has to face hypoxia and hydrogen sulphide in higher concentrations in its natural habitat.     

Marenzelleria cf. wireni (Polychaeta: Spionidae) from the Tay estuary. Metabolic response to severe hypoxia and hydrogen sulphide

The appearance of the spionid polychaete Marenzelleria spp. in the Tay estuary (Scotland) was first reported in 1984. Since estuaries are known as environments where abiotic conditions fluctuate widely, a laboratory study was conducted aimed to elucidate how this introduced species deals with low oxygen concentrations and exposure to hydrogen sulphide. Experimental evidence reveals that during severe hypoxia Marenzelleria cf. wireni switched to anaerobic pathways known from other marine invertebrates. Under the influence of sulphide (1 mmol l^-1) accumulation of anaerobic end products is more pronounced. It is assumed that in the presence of sulphide, due to its inhibition of aerobic respiration, the worms have to switch to anaerobic energy production directly after the onset of hypoxia. Seasonal differences in the metabolic reaction were found. It was shown that in specimens that were filled with gametes (winter), accumulation of end products of anaerobic metabolism was more pronounced than in immature worms (summer). The amount of succinate, an indicator for anaerobic energy production for instance, was nearly twice as high in the winter specimens. Since the amount of thiosulphate (a typical sulphide detoxification product) in the tissues is relatively low, it is suggested that Marenzelleria cf. wireni also accumulates other detoxification products. Sulphite, however, also known as a detoxification product, was found only in traces. Field measurements provide evidence that the potential of Marenzelleria cf. wireni to survive low oxygen and high sulphide concentrations is clearly higher than that normally needed in the Tay estuary.     

Food consumption of vendace Coregonus albula larvae in Lake Lentua, Finland

The daily ration of vendace larvae (mean TL 10·1–12·8 mm) was assessed after a time-intensive field survey in Lake Lentua. Ten larvae were sampled every 2 h throughout the 10-day sampling period and their alimentary tract contents analysed. The gut was cut into four quarters of equal length and each quarter was analysed separately. This procedure enabled the assessment of the proportions of different zooplankton taxa in the diet based on the least degraded first-quarter contents and to reconstruct the entire gut contents. Secondly, the gut passage time of ingested zooplankton was evaluated by comparing the degradation rate of zooplankton in the guts of experimentally fasted larvae to the degradation of zooplankton in the gradient from pharynx to anus among the larvae sampled from the lake. Finally, food consumption was calculated as zooplankton individuals and carbon biomass based on the two variables, gut passage time of constantly feeding larvae and reconstructed gut contents. The daily ration of vendace larvae varied between 14 and 450 zooplankters in the monitored period.     

Population dynamics, growth and production of the neozoon Marenzelleria cf. viridis (Verrill, 1873) (Polychaeta: Spionidae) in a coastal water of the southern Baltic Sea

The propagation of an immigrant from North America, viz. the spionid Marenzelleria cf. viridis in the Darss-Zingst Bodden (DZB) (southern Baltic Sea), was studied at three stations from March 1992 to December 1995. Highest mean abundances (over 28 000 ind. m^-2) and wet weights (400 g m^-2) were recorded at station 2 in 1994. The spionid also reached its highest dominances in terms of biomass (40 to 90%) at this station, which was selected for the population dynamics, growth and secondary production studies. The spionid has a life span of about 3 years, and many individuals achieved sexual maturity after one year. Their growth curve is steepest during the first year of life, during which the animals grow to a length of about 180 segments. However, growth depression was observed during the ripening of the gametes in April, May and June. Secondary production was in the region of 55 to 85 g AFDW m^-2 y^-1. Productivity (P/B) varied considerably from generation to generation, ranging between 0 and 4.8 with an average between 1.2 and 1.6.     

Gut fluorescence in herbivorous copepods: an attempt to justify the method

Recently the gut fluorescence technique has been critisized because of the possible degradation of chlorophyll into nonfluorescent derivatives during passage through copepod guts and changes of the gut passage time with food concentration. Here pigment budgets have been calculated in 6 experiments with Calanus finmarchicus CIV caught 2 km offshore of the Murmansk Marine Biological Institute (the Barents Sea, Dalnije Zelentsi) in September 1992. Copepods were fed with culture of Platymonas viridis at different concentrations. Gut pigment and ingestion rate increased with food concentration in a similar way. On average between 78% and 89% of ingested chlorophyll was recovered in the guts and faecal pellets. No trend for a greater loss of fluorescence at low food concentration than at high was observed. Pigment content of faecal pellets incubated in filtered seawater decreased by 20–30% in the first 7–12 h and by up to 60% in 48 h. The decline of pigment content was accompanied by a rapid bacterial growth (by a factor of 3 in 48 h). Gut passage time increased with decreasing food concentration (from 40 min at 9 µg pigm l^–1 to 64 min at 0.9 µg pigm l^–1). These results together with some data by other authors suggest that the gut fluorescence method can be used to estimate in situ grazing rate providing gut passage time is measured properly and there are no losses of faecal material. However, careful consideration should be given to the previous feeding history of copepods.     

The effect of seed morphology on the potential dispersal of aquatic macrophytes by the common carp (Cyprinus carpio)

1. The potential for seed dispersal by fish (ichthyochory) will vary among aquatic plants because of differences in seed size and morphology. 2. To examine how seed morphology influences the probability of dispersal by the common carp (Cyprinus carpio), we studied seed ingestion, retention time and subsequent egestion and germination of seeds of Sparganium emersum and Sagittaria sagittifolia, two aquatic plant species with similar sized but morphologically different seeds. 3. We compared dispersal probabilities between the two plant species, in which the probability of dispersal is assumed to be a function of the probabilities of seed ingestion, egestion and germination, and the dispersal distance is assumed to be a function of seed egestion rate over time. 4. We found that, although the soft seeds of S. sagittifolia had an approximately 1.5 times higher probability of being ingested by the carp than the hard seeds of S. emersum (83.15% ± 1.8% versus 56.16% ± 2.7%, respectively), the latter had an almost twofold higher probability of surviving the passage through the digestive tract (38.58% ± 2.7% versus 20.97% ± 1.5%, respectively). Patterns of seed egestion over time did not differ between the two plant species, despite the difference in seed morphology. Gut passage had a different effect on seed germination between plant species. Compared with non‐ingested controls, seeds of S. emersum showed a 12.6% increase in germination and a 2.1 day acceleration in germination rate, whereas seeds of S. sagittifolia displayed a 47.3% decrease and 5.1 day delay, respectively. 5. Our results suggest that seed morphology affects the dispersal probability and postdispersal establishment, but not the dispersal distance, of aquatic plants that are dispersed by fish.     

Reproductive success of the rotifer Brachionus calyciflorus feeding on ciliates and flagellates of different trophic modes

SUMMARY 1. The nutritional value of the bacterivorous ciliate Tetrahymena pyriformis and the algivorous ciliate Coleps sp., as well as the heterotrophic flagellate Chilomonas paramecium and the autotrophic flagellate Cryptomonas ovata , were investigated in population growth experiments using the rotifer B. calyciflorus . The two ciliates, both flagellates, which were of similar size, shape and mobility, were each offered as a sole diet and as a supplement to the alga Monoraphidium minutum , known to support reproduction of B. calyciflorus .
2. To further test nutritional differences between the prey organisms, prey selection experiments were conducted in which B. calyciflorus was able to select between the bacterivorous and algivorous ciliate, and between the heterotrophic and autotrophic flagellate.
3. The results demonstrated that both ciliates and the heterotrophic flagellate were not sufficient to support reproduction of B. calyciflorus when offered as a sole diet. They were, however, a good supplement to algal prey (except for the bacterivorous ciliate T. pyriformis ). In the prey selection experiments, B. calyciflorus positively selected for the algivorous Coleps sp. and the autotrophic C. ovata.
4. Overall, ciliates and heterotrophic flagellates may enhance survival of B. calyciflorus , but reproduction of the rotifer is likely to rely on algal prey. Both higher population growth of B. calyciflorus when fed the algivorous Coleps and the autotrophic Cryptomonas, along with their positive selection, give evidence for prey specific differences in nutrition, with algivorous or autotrophic prey species tending to be of higher nutritional value.     

The effect of phytoplankton and suspended sediment on the growth of Corbicula fluminea (Bivalvia)

Juvenile Corbicula fluminea Müller (1774) were cultured at 15.3 °C in the laboratory on eight combinations of suspended sediment and phytoplankton. Sediment concentrations were 2.6, 25, 50, and 150 mg l^–1. Chlorophyll a levels were 15.6 and 62.5 µg l^–1. Clam tissue growth was found to be independent of silt concentration but increased at the higher chlorophyll level (p < 0.05). The growth experiment was repeated at 24 °C with chlorophyll a concentrations of 18.9 and 112.6 µg l^–1. Growth was again greater at the higher phytoplankton level (p < 0.05). These results demonstrate that Asiatic clam populations are food-limited most of the growing season in the Northern and Western portions of California's eutrophic Sacramento-San Joaquin Delta where chlorophyll a levels average less than the lower of these values. Comparisons of clam growth in the laboratory and estuary support the food limitation hypothesis as at the higher food concentration laboratory tissue growth was 2.3 and 3.8 times greater during the high and low temperature evaluations than in the estuary.     

Zooplankton nutrition: recent progress and a reality check

Evidence suggests that marine and freshwater zooplankton generally experience food levels above subsistence values in terms of carbon. However, the quality of this food may be poor due to an insufficiency of other essential nutrients. In this review, we examine recent progress in three main areas of food quality research: (1) elemental (especially P) limitation, (2) digestion resistance, and (3) biochemical (especially fatty acids) limitation. We evaluate laboratory and field evidence in each of these areas, look at new evidence about the life history implications of the elemental limitation hypothesis, and suggest future avenues for research. From a rather large number of seemingly heterogeneous studies, a single consistent picture of food quality emerges: both P and essential fatty acids are predicted to be important dietary factors, but at different places and times. Nevertheless, despite an abundance of valuable laboratory studies, our knowledge of food quality limitation in the field is still poor.     

Sexual conflict over copula timing: a mathematical model and a test in the yellow dung fly

Sexual conflict over mating occurrence, timing, or durationis common in animals. This explains conspicuous female materejection behavior in many species, often involving shaking,fighting, and occasional forced copulations. We present a simplemodel that generates predictions about whether and when copulationoccurs in such conflict situations and how much female rejectionbehavior should be observed. Predictions depend on 2 underlyingparameters affecting female resistance and male persistence.We supply 2 qualitative tests of the model using the yellowdung fly Scathophaga stercoraria (Diptera: Scathophagidae).We manipulated adult age, body size (large and small), and adultfood availability (low and high), independently in males andfemales, staging replicate pairings of all treatment combinations.In agreement with predictions of our model, shaking durationfirst increased to a maximum at intermediate age, when the averagefemale copulated, and then decreased again. Contrary to expectation,body size did not affect copulation timing, female resistance,or male persistence. As predicted, adult food limitation delayedsexual maturity and hence prolonged female resistance, resultingin later copulations after more shaking. However, although foodlimitation equally delayed the increase in male persistencewith age, copulation also occurred later after more shaking,opposite to the model prediction. We conclude that shaking isdriven primarily by female age and male responses to it. Althoughfemale shaking can initially successfully deter males in S.stercoraria, this behavior is subtle and has apparently shiftedfunction from an effective means of mate choice to a signalof nonreceptivity, though its importance in nature remains unclear.     

Stoichiometric food quality and herbivore dynamics

Herbivores may grow with nutrient or energy limitation, depending on food abundance and the chemical composition of their food. We present a model that describes herbivore growth as a continuous function of two limiting factors. This function uses the synthesizing unit concept, has the hyperbolic Monod model as a limiting case, and has the same number of parameters as the Monod model coupled to Liebig's discontinuous minimum rule. We use the model to explore nutrient-limited herbivore growth in a closed system with algae, Daphnia and phosphorus as the limiting nutrient. Phosphorus in algae may substantially influence Daphnia growth. This influence changes over time and is most pronounced when algae and Daphnia populations fluctuate strongly. Relative to classic models that only consider food quantity as a determinant of Daphnia growth, our model shows richer dynamical behaviour. In addition to the standard positive equilibrium, which may be stable or unstable depending on nutrient availability, a new positive equilibrium may arise in our model when mortality rates are relatively high. This equilibrium is unstable and reduces the likelihood of long-term persistence of Daphnia in the system.     

Sex-limited expression of ornamental feathers in birds

Extravagant secondary sexual characters show sexual size dimorphismin some species but are completely sex limited in others. Sexualornamentation has been hypothesized to benefit mainly malesthrough sexual selection, but the costs of secondary sexualcharacters initially would be experienced by both sexes. Theevolution of sexual size dimorphism of ornaments and, eventually,the complete sex-limited expression of these characters, willdepend on the effects of sexual and natural selection on thetwo sexes. A phylogenetic analysis controlling for similaritiesdue to common ancestry of 60 independent evolutionary originsof feather ornamentation in birds was used to investigate ecologicalfactors correlated with sexual size dimorphism and sex-limitedexpression of secondary sexual characters. When the size ofan ornament is large relative to body size, the trait willbe particularly costly for females, resulting in selectionfor increased sexual size dimorphism of the ornament. Indeed,sexual size dimorphism of ornaments was positively relatedto the relative size of male ornaments but was unrelated torelative size of female ornaments. Species with polygynousand lekking mating systems with little or no male parentalcare (in particular nest building and incubation) demonstratedsex-limited expression of ornaments as compared to monogamousspecies. Species with no food provisioning of offspring by themale showed a trend for increased sexual size dimorphism ofornaments. Therefore, large natural selection costs duringreproduction imposed by the expression of secondary sexualcharacters are related to the evolution of sexual size dimorphismof ornaments and eventually their complete loss from females.