Genetic variability in bloodworm (Glycera dibranchiata) populations in the Gulf of Maine

Genetic structure of eight Northwest Atlantic populations of the marine polychaeteGlycera dibranchiata Ehlers was examined with starch gel electrophoresis. Samples were collected during summer and fall 1981, and seven polymorphic and four monomorphic loci were consistently scored. Average heterozygosity (0.126) and percent polymorphic loci (59.3) were comparable to the averages reported for marine invertebrates. Minimum genetic distances between populations ranged from 0.003 to 0.093, levels typically associated with local populations of the same species in other taxa. Based on these data, inter- and intra-estuarine migration and gene flow appear to be low. Only two populations, separated by 13 km along the same river in New Brunswick, Canada, were not genetically different from each other. These findings may have relevance for management strategies in bloodworms.     

Reproductive behaviour ofNereis diversicolor (Annelida: Polychaeta)

Nereis diversicolor O. F. Müller, collected on the tidal flats of the Jadebusen (North Sea, FRG) in autumn 1987 and 1988, was exposed to different temperatures in the laboratory. Results indicated that maturation was induced by temperatures above 6°C; spawning in early spring was synchronized by raising temperatures after a period of low temperatures in winter, and occurred mainly at new and full moon. During reproduction the female stays inside the burrow; the male releases sperm in front of it; the resultant larvae remain in the tube for 10 to 14 d protected by the female.     

Secondary production ofPrionospio caspersi (Annelida: Polychaeta: Spionidae)

The annual somatic production ofPrionospio caspersi Laubier was estimated, between July 1986 and 1987, at a sandy shallow-bottom station in the area facing the Po River Delta, Italy, where this species is one of the most abundant. Primary recruitment occurs in summer, and maturation of gametes the following spring. Body sizes were estimated from the surface area of the anterior end of worms, projected onto a monitor and recorded using a digitizer tablet. Measurements, to the tenth setiger, were correlated with total length, total body surface area and AFDW. The annual production was 8.06 g m^–2 yr^–1 (AFDW) and the production:biomass (P:B) ratio 4.09. The results are discussed in relation to some physical and chemical features of the environment, stressing differences with the life cycle of other coexisting populations.     

An investigation on the anaerobic metabolism of Nephtys hombergii (Annelida: Polychaeta)

In Nephtys hombergii Savigny (Annelida: Polychaeta: Errantia), succinate and less pronounced L-alanine are accumulated as endproducts during an initial phase of anaerobiosis. In this phase aspartate is utilized as substrate for anaerobic energy production in addition to glycogen. Prolonged anaerobiosis results in the formation of propionate and acetate which, to a large extent, are excreted into the water. The concentrations of aspartate and succinate were found to remain unchanged down to a Pw_{O 2}(oxygen partial pressure in incubation water) of approximately 20 torr, indicating a fully aerobic metabolism. At a Pw_{O 2}of 15 torr characteristic changes can be observed: energy production becomes partially anaerobic. At a Pw_{O 2}of 7 torr the level of aspartate is largely reduced, but the accumulation of succinate is still significantly less than at anoxia. [In the habitat of N. hombergii, the intertidal mud flats, a Pw_{O 2}in the range from 0 torr (Pamatmat, 1968) to 12 torr (Wells and Warren, 1975) was measured.] The activities of some important enzymes involved in anaerobic energy production (among them pyruvate kinase, phosphoenolpyruvate carboxykinase and lactate dehydrogenase) were measured. Although N. hombergii has a high glycolytic capacity, no lactate dehydrogenase was detected. Instead strombine and alanopine dehydrogenases were present in comparatively high activities.     

Cysteine-glutathione disulfide, the sperm-release pheromone of the marine polychaete Nereis succinea (Annelida: Polychaeta)

Summary. In the marine polychaete Nereis succinea (Frey & Leuckart 1847) a sex pheromone was isolated from the coelomic fluid of sexually mature females and identified by NMR studies and independent synthesis. This pheromone is released by the females during reproduction together with eggs and coelomic fluid into the free water column and induces sperm release of surrounding males. Its structure was ascertained as L-cysteine-glutathione disulfide. It exhibited a response threshold of 0.6 · 10⁻⁷ M. Received 15 July 1997; accepted 25 November 1997.     

Nachweis der epitokie als fortpflanzungsmodus des polychaeten Eunice siciliensis (Polychaeta: Annelida)

Eunice siciliensis (Grube)¹ is a sedentary polychaete with separate sexes; its germ cells develop only in the posterior part of the body. In March and April, females with whitish or even dark-green oocytes, males with only few spermatogonia, and individuals with no germ cells at all were discovered in the “coralligène” of the Banyuls region (European Mediterranean Sea). In august, besides immature and sexually non-differentiated worms, mature individuals of both sexes were captured: females with dark bluish-green oocytes of diameters up to 250μm, and males with milky genital segments containing spermatozoa. Soon after isolation from the substrate, these individuals performed characteristic movements with their posterior body parts, which then autotomized, and released ova and spermatozoa. Evidence is given that the anterior, atokous body parts survive in their tubes in the “coralligène” after separation from the epitokous portions, and regenerate new genetal segments. The development of gametes observed in these regenerated segments indicates the ability of E. siciliensis to reproduce more than once during its lifetime. Not only caudal, but also prostomial regeneration was observed; the formation of a (morphologically different) secondary prostomium appears to be essential for any development of germ cells in worm fragments. E. siciliensis exhibits surprising similarities in habits and in formation of an epitokous form with Eunice viridis (Gray), the “palolo” worm of the South Pacific Ocean, but nevertheless, lacks the prominent criteria of this famous species: the ventral eye spots on each genital segment in males and females, and the paired areas with brown pigmentation on the ventral side of each epitokous segment in the males.     

Consumption of diatoms and diatom filtrates by the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta)

Rates of organic uptakes of three live diatoms (Nitzschia acicularis, Nitzschia sp. andNavicula incerta), and of three corresponding filtrates by the deposit-feeding polychaeteEupolymnia nebulosa (Montagu) were measured under similar experimental conditions. Worms used during this study were collected by SCUBA diving at Port-Vendres in shallow water (7 m deep) during the summer of 1986. Uptake rates of live diatoms were affected both by length of the experiments and by the nature of the food offered. The highest rate of uptake (11.8 10^–4 mg algal ash-free dry wt mg^–1 worm dry wt h^–1) was recorded during a short-term experiment (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate (1.1 10^–4 mg algal ash-free dry wt mg^–1 worm dry wt h^–1) was recorded during a long-term experiment (48 h) with the largest diatom (Nitzschia acicularis). Filtrates ofN. acicularis were more readily utilized than those ofNitzschia sp. andNavicula incerta. Because of differences in uptake rates of algal filtrates as a function of species, it is not possible to evaluate the bias due to interaction with dissolved substances in experimental studies assessing ingestion of live benthic diatoms byE. nebulosa.     

Effects of temperature on oocyte growth in the Mediterranean terebellid Eupolymnia nuebulosa (Annelida: Polychaeta)

Eupolymnia nebulosa (Montagu) is a widely distributed terebellid polychaete that builds its tubes on the coastal shelf in areas with mixed soft and hard bottoms. From a long-term survey in the Bay of Banyuls, France (NW Mediterranean), we found an advancement of the timing of the spawning period coincident with a delayed breakdown of the thermocline. We postulate that persistent high temperatures can influence gamete development by stimulating oocyte growth, resulting in earlier spawning. During 1992 and 1993, we used a between-individuals experimental approach to assess the possible effect of temperature on oocyte growth based on: (1) determination of the growing fraction of the oocyte population (i.e. oocyte net growth); (2) identification of differences in oocyte growth-rate among females; (3) comparison of non-significantly different size-distributions of the growing oocyte fraction (net oocyte size-distributions) at the beginning of the experiments with those at the end. No effect of temperature on oocyte growth was detectable at the population level, but a positive individual response to prolonged high temperature was evident. Thus, the lack of a significant response by the population to prolonged high temperature does not imply a lack of individual response. We propose a model of oocyte-growth dynamics based on temperature that incorporates previous observations of extended oogenesis and oocyte growth during periods of both increasing and decreasing temperature and on the scattered pattern of oocyte size-distributions at the onset of spawning. Received: 4 November 1996 / Accepted: 18 January 1997     

Feeding responses of the tentaculate deposit-feederEupolymnia nebulosa (Annelida: Polychaeta): Influence of sexual maturity

Feeding responses of the tentaculate depositfeeding polychaeteEupolymnia nebulosa (Montagu) were studied by measuring rates of uptake of three different¹⁴C-labelled diatoms (unialgal cultures ofNavicula incerta Grunow,Nitzschia acicularis Wm Smith, andNitzschia sp.). Worms used during this study were collected in the harbor of Port-Vendre (Western Mediterranean) during August 1986 (immature worms) and December 1987 (mature worms). Uptake rates were affected both by the length of the experiments and by the nature of the food offered. The highest rate of uptake (2.98 10^–4 mg ashfree dry wt of algae mg^–1 dry wt of worms h^–1) was obtained during short-term experiments (4 h) with the smallest diatom (Nitzschia sp.). The lowest rate of uptake (0.21 10^–4 mg ash-free dry wt of algae mg^–1 dry wt of worms h^–1) was also obtained withNitzschia sp., but for a long-term (48 h) experiment. There was no significant difference between rates of uptake of immature and mature worms.     

Measurement of the adenylate energy charge in Nereis diversicolor and Nephtys sp. (Polychaeta: Annelida):

ATP-content and adenylate energy charge (AEC) ratios were determined in two polychaete species (Neries diversicolor and Nephytys sp.), sampled in ten stations along the heavily polluted Western Scheldt estuary (N. Belgium, S. Holland). The samples were taken between 27 December 1982 and 6 January 1983. Nereis diversicolor was also sampled in an unpolluted brackish water pond, and subjected to artificial stress by drying the organisms on filter paper. Adenine nucleotide levels were determined using the firefly bioluminescence reaction. The light emission was measured with the integration mode. As the Constant Light Signal reagent used contained AMP and probably ADP itself, both myokinase and pyruvatekinase had to be inactivated before the assay. The use of pepsin in acidic conditions to denaturate these enzymes is discussed. For both N. diversicolor and Nephtys sp., a significant relation between ATP-content and fresh weight is found in most samples. Within each species, ATP-content per g fresh weight is dependent on the sampling method, but it is not significantly different between stations. It is higher in Nephtys sp. from the W. Scheldt (130. 8 10^-5 g ATP/g fresh weight) than in N. diversicolor from the Dievengat (83.9 10^-5 g ATP/g fresh weight). This value in turn is significantly higher than that for N. diversicolor in the W. Scheldt (64.0 10^-5 g ATP/g fresh weight). In the drying experiment, AEC gradually decreases with increasing drying time. In the field, AEC is generally high and not significantly different between the three populations. AEC ratios also do not differ between stations within each population. It is argued that field monitoring of AEC in invertebrates is not valuable as a pollution assessment tool for two reasons. In polluted stations, only pollutant-resistant species will be found, and there is evidence that the maintenance of a stable population is impossible when the individuals constantly have low AECs.     

Isozymes and general protein patterns as taxonomic markers in the taxon Nephtyidae (Annelida Polychaeta)

Seven Nephtys species and one species of the genus Aglaophamus (collected from different European tidal and subtidal locations between 1989 and 1991) were compared with respect to six isozyme systems (-amylase, esterase, hexokinase, malate dehydrogenase, malic enzyme, phosphoglucoseisomerase) as well as with respect to general protein patterns with non-specific staining. The proteins were obtained from the tissue (proboscis, individual segments) of single individuals, the species of which had previously been accurately determined, and were separated by isoelectric focusing (IEF) in polyacrylamide gels. The enzymes were identified by their specific catalytic activities, and the general proteins were visualized by silver staining. All the isozymes proved to be monomorphic within each of the various polychaete populations. With a single exception, it was impossible to distinguish geographically separated populations of the same species, because the band patterns were completely consistent within the species. The individual enzymes varied in their suitability for species differentiation. Only with respect to esterase and -amylase could all species be distinguished; the other enzymes studied were identical in morphologically similar species. In contrast, each species could be identified by its general protein pattern, although some species differed from others in the position of a few bands only. Individuals of N. hombergii, N. caeca and N. cirrosa with abnormally shaped parapodia, as well as juvenile nephtyids, could be unequivocally assigned to their respective species by IEF. Only in the case of N. longosetosa were two morphological variants found to differ in their electrophoretic characteristics; one of them is interpreted as a new species, not previously described.     

Dietary effects on oocyte yolk-composition inCapitella sp. I (Annelida: Polychaeta): Fatty acids and sterols

In 1988, immatureCapitella sp. I (initial biomass 80µg dry wt) were raised on four diets: Gerber cereal, TetraMin fish food,Ulva sp., and benthic diatoms. After 2 wk of culturing different populations on these diets, eggs were dissected from gravid females, frozen and analyzed for fatty acid and sterol composition. Eggs produced by worms on different food types were discriminated by fatty acid composition in a principal component analysis (PCA), with 18:1⁹, 18:2, 20:5 and 20:4 fatty acids showing the greatest differences between experimental groups. The sterol profiles of all eggs were dominated by cholesterol (cholest-5-en-3-ol) and cholest-5,24-dien-3-ol (>60%). A PCA of egg sterols discriminated between adult diets with cholesterol, 23,24-dimethyl-cholesta-5-en-3-ol, cholest-5,24-dien-3-ol and a C-29 stenol showing the greatest differences. In field populations ofCapitella sp. I, oocytes produced at different times of the year may have different levels of lipids depending on their dietary availability during vitellogenesis. Variations in oocyte composition may influence larval growth and development and thus have an impact on population dynamics. Alternatively, variations in fatty acid and sterol composition of oocytes may allow us to identify the food resources utilized by this species.     

The relation between spontaneous activity cycles and diurnal rhythms of metabolism in the polychaetous annelid Glycera dibranchiata

The oxygen consumption rate of the bloodworm Glycera dibranchiata Ehlers was measured 4 times each day. Aerobic metabolism during daylight hours is significantly greater than at night, regardless of the phase of the spontaneous activity cycle. There is no reason to suppose that increased metabolism reflects an overall increase in motor activity.Supported by National Science Foundation research grant GB-6884.     

Utilization of a fumigated sediment by two benthic deposit-feeders:Abra alba (Mollusca: Bivalvia) andEupolymnia nebulosa (Annelida: Polychaeta)

The present study tested the utilization of dead microbial biomass by two benthic deposit-feeders:Abra alba (Wood) (Mollusca: Bivalvia) andEupolymnia nebulosa (Montagu) (Annelida: Polychaeta). Clams were collected in the Canet lagoon during spring 1989. Worms were collected in the Port-Vendres harbour during spring 1989. The¹⁴C-labelled (glutamic acid, 24 h) sediment used during the study was sterilized with 1% chloroform, washed with sterile seawater, and dried (60°C; 48 h). This sterilisation procedure, called fumigation is the least harmful to the sediment (Novitsky 1986). Both clams and worms were incubated in the presence of the fumigated sediment for 5, 10, 20, and 50 h. At the end of each experiment we recorded the radioactivity in four compartments: (1) sediment, (2) dissolved organic matter (DOM), (3) CO₂, and (4) animals. The radioactivity of the sediment was subdivided into five fractions: (i) soluble in 2N HCl, (ii) soluble in hot 5% trichloroacetic acid (TCA), (iii) soluble in 1N NaOH, (iv) soluble in hot 6N HCl, (v) residual (after combustion in a Leco carbon analyser). In the first set of experiments, after 20 h of incubation, 5.4 and 4.7% of the total radioactivity was taken up by clams and worms, respectively. However, a model revealed that this uptake could have been correlated with the release of radiolabelled DOM (33% of total radioactivity during the first 5 h). In order to test this assumption, we used the same protocol with three additional washes of the fumigated sediment. This resulted in a significantly lower uptake by the clams (1.9% of the total radioactivity byt = 50 h), whereas the worms exhibited an uptake similar to that in the initial experiment (5.1% of total radioactivity byt = 50 h). These results underline the importance of considering interactions with DOM when applying radiotracer techniques to the study of benthic food chains. The average ingestion rates of fumigated sediment byA. alba andE. nebulosa were 5.2 10^–2 mg sediment dry wt mg^–1 clam h^–1 and 3.5 10^–2 mg sediment dry wt mg^–1 worm h^–1, respectively, which is comparable to previous data reported for other deposit-feeding bivalves and polychaetes feeding on natural sediment or detritus. The low radioactivity recorded for CO₂ together with the similarity of the changes in the partitioning of the radioactivity within the sediment between control experiments and experiments carried out in the presence of clams or worms suggest low assimilation efficiencies. Therefore, the present study supports the fact that dead microbial biomass does not constitute an important food source for benthic deposit-feeders.     

Effect of food type and ration on growth of juvenile Capitella sp. I (Annelida: Polychaeta): macro- and micronutrients

Growth rates of juvenile Capitella sp. I were determined on different rations of six food types: Gerber's mixed cereal, TetraMin fish flakes, benthic diatoms, Ulva sp., spring detritus, and summer detritus. A simple growth model estimated maximum growth rate and maintenance ration for each food. There were differences in the growth response among foods relative to nitrogen content. As juveniles increased in size, differences in growth between foods became more pronounced. For all juveniles, growth rates were correlated with levels of the amino acids histidine, phenylalanine, threonine, and valine, and the polyunsaturated fatty acid 20:5w3; correlations with histidine and phenylalanine levels were the most significant. Regressions of growth rates as a function of these two amino acids suggest a daily maintenance ration of 300 pg histidine and phenylalanine mg^-1 nitrogen biomass. Juvenile worms grew on spring detritus but not on summer detritus, indicating the probable importance of micronutrients (polyunsaturated fatty acids, amino acids) for growth of juvenile Capitella sp. I in the field.     

Uptake and loss of radioactive cadmium by the sea-skater Halobates robustus (Heteroptera: Gerridae)

Sea-skaters, Halobates robustus Barber, caught around the Galápagos Islands, were exposed over a period of 3 days to radioactive cadmium dissolved in seawater. Although ^115mCd (metastable ¹¹⁵Cd) was accumulated by the insects, they did not concentrate it above the seawater level. The experiments clearly showed that cadmium is taken up into the body, not merely adsorbed onto surfaces. The uptake of Cd followed a biphasic time course. When sea-skaters labelled in this way were transferred back into unlabelled seawater, the rate of loss of Cd was likewise biphasic. During uptake and loss, the biological half-life times for the first compartments were almost identical (0.64 and 0.79 h), indicating that the same compartment might be involved. On the other hand, the biological half-life times for the second compartments were different (1.7 days during uptake, 19.5 days during loss), indicating that the site or chemical form of bound Cd in the sea-skater had changed. For an adult female, weighing about 8.5 mg wet weight, the drinking rate of seawater was estimated at 2.5 l day^-1, equivalent to 29% body weight.     

Gametogenesis in Harmothoe imbricata (Polychaeta: Polynoidae)

The processes of gametogenesis in Harmothoe imbricata are described and illustrated. For most of their development the gametes are attached to the walls of segmental blood vessels. In males, only meiosis and sperm differentiation occur while the cells are free in the coelom. In females, oocytes are only released into the coelom briefly, after vitellogenesis has been completed. H. imbricata thus differs from most polychaete species so far described — where in males the spermatogonia divide mitotically to form rosettes after they have been released into the coelom, and in females the whole of vitellogenesis occurs after release of oocytes into the coelom. The oocytes of H. imbricata enter the first metaphase of meiosis before they are shed at spawning. The elongated head of the sperm in H. imbricata is atypical of polychaetes in general.     

Etude expérimentale du comportement nuptial de Platynereis dumerilii (Annelida: Polychaeta): chémoréception, émission des produits génitaux

In the heteronereis form of Platynereis dumerilii, the roles of the swelling of the parapodial cirri of the male or female first segments, of the crenellations of the dorsal parapodial cirri of the male posterior region and those of the male pygidial papilli have been ascertained experimentally with respect to nuptial behaviour. The nuptial dance and the release of the genital products are provoked by chemical stimulating discharges from the partner of the opposite sex. The chemical stimuli act on the swelled or crenellated parapodial cirri, probably through the bipolar chemoreceptor neurons which are numerous on these organs. The excitation is transmitted to the cerebral ganglia by way of the ventral cord; it is translated into motor impulses which act upon the musculature concerned in the nuptial dance and in the release of genital products. Accordingly, the swelling and the crenellations of the parapodial cirri appear to contain chemoreceptors. The specificity of this chemoreception, in regard to the exciting substance emitted by the opposite sex, is probably related to the presence of a protein (Boilly-Marer, 1966a, b) elaborated during heteronereidation by epidermal cells adjacent to the dendritic extensions of the chemoreceptor cells. The pygidial papilli appear to insure sperm evacuation.     

Ovary structure and oogenesis in Cirratulus cirratus (Polychaeta: Cirratulidae)

Oogenesis in polychaetes has, in most species, two main phases: a coelomic phase during which vitellogenesis occurs, and an ovarian phase. Details of the latter are known in very few species. Efficient reproduction requires coordination between these two phases of oocyte differentiation. The ovaries of Cirratulus cirratus (O. F. Müller) occur 2/segment throughout the fertile region of the body; 5 main stages of oocyte differentiation can be recognised on them: stem cells, oogonia, premeiotic oocytes, early oocytes and terminal oocytes. A quantitative study of the composition of the ovary throughout the reproductive cycle shows that the ovary size is approximately constant, but that the percentage of cells in the terminal oocyte stage is reduced to zero prior to spawning. This stage in oocyte differentiation is therefore a critical one, at which oogenesis may be arrested. The ovary alternates between a proliferative phase when the terminal oocytes are present, and a non-proliferative phase when they are absent. The overall rate of germ cell production for each ovary is low, less than 1 cell/ovary/day, and the, ovary tissue turnover time is greater than 2 years. A qualitative model describing the production of oocytes is presented, and the implications of the low rate of germ cell production are discussed. A low rate of germ cell proliferation, as in Cirratulus cirratus, implies that the rate of oocyte output into the coelom will be controlled by factors influencing the later stages of ovarian oocyte differentiation.