Nereis diversicolor effect on the stability of cohesive intertidal sediments

The effect of the polychaete Nereis diversicolor on the stability of natural cohesive sediments was investigated in the laboratory. Three densities (450, 600 and 1200 ind m⁻²) of N. diversicolor were used. Sediment shear strength was measured using a cone penetrometer. Sediment erodability was assessed using an annular flume (current velocities from 5 to 55 cm s⁻¹) in which flow velocity was increased incrementally, and water sampled to quantify suspended material in order to derive critical erosion velocity and erosion rates. At low current velocities ( <25 cm s⁻¹), we found N. diversicolor to have a stabilising effect, reflected by an increase of up to 20% in the critical erosion velocity. This is related to an enhancement of ~50% in shear strength, due probably to gallery building activities, responsible for the promotion of lateral compaction, an increase in the area of the sediment–water interface, and enhanced microphytobenthos production. Once the sediment began to erode, the stabilising effect of N. diversicolor reverses, leading to an increase of up to 40% in eroded matter due to compaction, which resulted in the erosion of larger aggregates. The balance between the effect of N. diversicolor on herbivory and microphytobenthos production due to the presence of galleries is discussed. Our results indicate that neither chlorophyll a, nor shear strength nor critical erosion velocity are good indicators of erodability. This underlines the need to include biogeochemical processes in any realistic sediment transport model.     

Remedial habitat creation: does Nereis diversicolor play a confounding role in the colonisation and establishment of the pioneering saltmarsh plant, Spartina anglica?

Increasing concerns over global warming and expected sea level rises have led to the adoption of new coastal management strategies around the south-east coast of England. This paper explores the role played by the estuarine invertebrate Nereis diversicolor in limiting the colonisation and establishment of the invasive pioneering salt marsh plant, Spartina anglica. The biology of N. diversicolor is briefly reviewed and data from field experiments are presented demonstrating significant negative effects of worm abundance on transplanted S. anglica biomass. Laboratory-based experiments demonstrated significant negative effects of N. diversicolor abundance on the survival of S. anglica seeds transplanted to sediment cores. The importance of estuarine invertebrates in engineering the mudflat habitat may confound the foreseen ecosystem services and function provided by saltmarsh management schemes. Received: 15 February 1999 / Received in revised form: 4 June 1999 / Accepted: 11 June 1999     

The larval nephridia of the brackish-water polychaete,Nereis diversicolor

The larval nephridia of the brackish-water polychaete Nereis diversicolor are described for the first time, and have been studied to determine if their times of development and structural characteristics are consistent with a role in the osmotic regulation of the larva. As shown in serial paraffin sections and by interference-contrast optics, the nephridia of the three-setiger larva consist of a single pair of very large metanephridia, arising in the 3rd larval setiger, but with their elongated terminal ducts and coiled ciliated tubules pushed forward into the 2nd setiger; their open metanephrostomes and anterior anchoring filaments lie dorsal to the 2nd set of setae. In contrast, the definitive or juvenile metanephridia, arising in the 4th and subsequently formed setigerous segments, have short terminal ducts and coiled ciliated tubules confined to the segments on which their external nephropores open; their nephrostomes are ventrally located and open into the rear of the next anterior segment. These findings are in contrast to the claims of Edouard Meyer (1887), who described two pairs of closed protonephridia in the 2nd and 3rd larval setigers of Perinereis cultrifera. Although it is not excluded that the single larval pair of metanephridia of N. diversicolor may arise as protonephridia, Meyer's claim of two pairs of larval protonephridia was an observational error. The larval nephridia of the marine Platynereis dumerilii resemble in form, but are considerably smaller than, those of N. diversicolor. It is concluded that the hypertrophied pair of larval metanephridia of N. diversicolor is an evolutionary adaptation to existence in habitats of low and unpredictably varying salinity. Their development occurs irrespective of the prevailing salinity; hence, it must be genetically determined.     

Assessment of the influence of confounding factors (weight,salinity) on the response of biomarkers in the estuarine polychaete Nereis diversicolor

The influences of salinity and body size on biochemical (activities of glutathione-S-transferase, lactate dehydrogenase (LDH), acetylcholinesterase and digestive enzymes amylase and CMCase), physiological (feeding and egestion rates, energy reserves) and behavioural (burrowing speed) biomarkers were examined in the infaunal polychaete Nereis diversicolor. Only a few biomarkers were affected, including increased egestion rate and activities of CMCase and LDH at higher salinity, and higher egestion rate in larger worms. These findings reinforce the status of N. diversicolor as a robust sentinel species for estuaries which are environments that are particularly productive but also particularly at risk.     

Bacterial expression and characterization of molluscan IDO-like myoglobin

The indoleamine 2,3-dioxygenase (IDO)-like myoglobin (Mb) is a unique type of Mb isolated from the buccal mass of several archgastropod species. Here, we expressed Sulculus diversicolor IDO-like Mb as a GST-fusion protein in bacteria. The visible spectrum of GST-fusion IDO-like Mb shows characteristic α- and β-peaks, indicating that it binds oxygen. To identify residues important in heme and oxygen binding, we constructed site-directed mutants. We initially replaced each of the 7 histidines of S. diversicolor IDO-like Mb with alanine. The spectra of three mutants (H74A, H288A, and H332A) revealed a remarkable loss of absorbance around 414 nm, indicating that they cannot bind heme. His⁷⁴, His²⁸⁸, and His³³² were also replaced by arginine or tyrosine. Neither H332R nor H332Y contains heme, suggesting that His³³² is the proximal ligand of IDO-like Mb. In contrast, both H74R and H288Y mutants were isolated in the heme-binding oxy-form. The autoxidation rates of these two mutants showed that they can bind oxygen as stably as wild-type. His⁷⁴ and His²⁸⁸ might be partially associated with heme-binding, but do not act as the distal ligand. The S. diversicolor IDO-like Mb seems to stably bind oxygen in a different manner from normal myoglobins.     

A review of the biology, ecology and potential use of the common ragworm Hediste diversicolor (O.F. Müller) (Annelida: Polychaeta)

The common ragworm Hediste diversicolor inhabits the shallow marine and brackish waters in the North Temperate Zone of the Atlantic. This infaunal species builds U or Y-shaped burrows in the soft sediments. H. diversicolor is gonochoristic and remains atokous throughout its life. Due to the facility of harvesting of individuals by hand and their maintenance, this species represents a good biological material for experimental studies. In consequence, the common ragworm served as a model for endocrine control of reproduction in Polychaeta and played a fundamental role in the study of immune defence in Polychaeta.Life history characteristics (longevity, spawning season, feeding tactics and growth) and population dynamics (sex ratio, density and biomass) vary greatly according to geographical location of the populations. In addition, inter-population morphological, biochemical and physiological differences have been noticed in individuals from different areas and different environmental conditions in the same areas and could be related to the limited dispersal capacity of the species. H. diversicolor interacts with its environment. The construction of burrows by individuals increases the sediment–water interface. When they ventilate their burrows, individuals verticalize oxic zones into the sediment and promote microbial and meiofaunal growth alongside their burrows. Moreover, the common ragworm is a species of commercial interest because of its use as bait in recreational fishing and as food in aquaculture. In order to alleviate environmental pressure caused by excessive demand over optimal sustainable yield of bait fisheries, intensive aquaculture has been proposed. The common ragworm could be also used as biomonitor of pollution in estuarine ecosystems.     

The Fate of [3H]Ecdysone in Three Species of Annelids

Summary

The metabolism of [³H]ecdysone was examined in 3 species of annelids: the bloodworm, Tubifex vulgaris (a freshwater oligochaete), the earthworm, Lumbricus terrestris (a terrestrial oligochaete) and the ragworm, Nereis divtrsicolor (a marine polychaete). One of these species, N. diversicolor, metabolised injected [³H]ecdysone into compounds which co-chromatographed on both reversed-phase and adsorption HPLC with authentic 20-hydroxyecdysone, 26-hydroxyecdysone and 20,26-dihydroxyecdysone, thus demonstrating the occurrence of 20-hydroxylation and 26-hydroxylation capability in the Annelida. Furthermore, [³H]ecdysonoic acid was also formed and excreted by N. diversicolor, suggesting that 26-oic acid formation is involved in ecdysteroid inactivation in this species. Other, as yet unidentified, radioactive metabolites were also excreted by N. diversicolor. Several metabolites of [³H]ecdysone were also detected in the other 2 species examined, T. vulgaris and L. terrestris.     

Macrofaunal community bioturbation along an estuarine gradient

A multidisciplinary study of the impact of bioturbation on sediment dynamics has been underway for some time at the Plymouth Marine Laboratory. The current programme has been founded upon the careful selection of six sites in the River Tamar, representative of important combinations of physical and biological variables. The paper presents preliminary results illustrating the contrasting importance of the speciesNereis diversicolor andNephtys hombergi as agents of bioturbation, given their different distributions across the six sites. Thus bioirrigation byN. diversicolor increases up the estuary and is greatest in the region of high suspended bed-load due to tidal pumping, where the consequences for chemical exchange processes between sediments and the water column may be most important. The sediment mixing effects ofN. hombergi are likely to be greater towards the seaward end of the estuary but ultimately the particular sediment types and the macrofaunal communities they support dictate the level of bioturbation in ways that do not necessarily relate to simple axial gradients along the estuary.     

Evaluation of protein extraction protocols for 2DE in marine ecotoxicoproteomics

In ecotoxicoproteomics, an accurate and reproducible extraction of proteins is a critical step for 2DE analysis and further protein identification using MS. The criteria for the assessment of protein extraction quality include protein yield, protein spots resolved in a 2DE gel, matched protein spots in replicate gels, reproducibility, and compatibility with MS. In this work, we evaluated three protein extraction systems, straightforward lysis buffer, trichloroacetic acid–acetone, and TRIzol reagent with some modifications, for the protein extraction from three animal species including mussel Mytilus galloprovincialis, flounder Paralichthys olivaceus, and polychaete Nereis diversicolor used in marine ecotoxicology. Our results indicated that these methods could extract significantly different protein profiles. The method using TRIzol reagent resulted in the most matched protein spots resolved in four replicate 2DE gels and highest reproducibilities for the gill of M. galloprovincialis and liver of P. olivaceus. However, a modified trichloroacetic acid–acetone solvent system was best for the whole soft tissue of N. diversicolor. This work provides the fundamental information of the extraction quality of protein extraction protocols from different marine animals, which may facilitate the selection of a suitable protein extraction protocol for ecotoxicoproteomics.     

Phylogeography and cryptic introduction of the ragworm Hediste diversicolor (Annelida,Nereididae) in the Northwest Atlantic

Many benthic marine invertebrates show striking range disjunctions across broad spatial scales. Without direct evidence for endemism or introduction, these species remain cryptogenic. The common ragworm Hediste diversicolor plays a pivotal role in sedimentary littoral ecosystems of the North Atlantic as an abundant prey item and ecosystem engineer, but exhibits a restricted dispersal capacity that may limit connectivity at both evolutionary and ecological time scales. In Europe, H. diversicolor is subdivided into cryptic taxa and genetic lineages whose distributions have been modified by recent invasions. Its origin in the northwest Atlantic has not been adequately addressed. To trace the age and origin of North American ragworm populations, we analyzed mtDNA sequence data (COI) from the Gulf of Maine and Bay of Fundy (n=73 individuals) and compared our findings with published data from the northeast Atlantic. Our results together with previous data indicate that two species of the H. diversicolor complex have independently colonized the northwest Atlantic at least three different times, resulting in two distinct conspecific assemblages in the Bay of Fundy and Gulf of Maine (respectively) that are different from the species found in the Gulf of St. Lawrence. North American populations had significantly lower genetic diversity compared with populations in the northeast Atlantic, and based on patterns of shared identity, populations in the Bay of Fundy originated from the Baltic Sea and North Sea. Populations from the Gulf of Maine were phylogenetically distinct and most likely originated from unsampled European populations. Analyses of the North American populations revealed patterns of post‐colonization gene flow among populations within the Gulf of Maine and Bay of Fundy. However, we failed to detect shared haplotypes between the two regions, and this pattern of complete isolation corroborates a strong phylogeographic break observed in other species.     

Herbivores and the Spatial Distributions of the Phytoplankton. I. The Plankton Market

Planktonic herbivores forage on phytoplankton cells which were produced at some earlier time and at a distant place. A steady state, one dimension model of the birth, death and eddy diffusion of phytoplankton illustrates phytoplankton smooth both temporal and spatial variations as they are transported. When the eddy diffusivity is k and the phytoplankton doubling time is b, the average distance traveled by a phytoplankton cell is (kb)^1/2 and the average time spent doing so is b. Only those spatial variations with wavelengths greater than 2π(kb)^1/2 and temporal variations with periods greater than 2πb will be observed in the phytoplankton distributions at more than half their original amplitude. Both k and b control the length scale of phytoplankton distributions in a cartesian coordinate system. Planktonic herbivores view the phytoplankton from a diffusing coordinate system in which the spatial scales of the phytoplankton distribution are transformed into time scales.     

Long‐term allelopathic control of phytoplankton by the submerged macrophyte Elodea nuttallii

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Accumulation of americium 243 in selected brackish and marine invertebrates

As an initial step in a programme designed to investigate factors which are of importance in affecting the behaviour of actinides towards certain invertebrates found in estuarine and marine environments, laboratory procedures have been developed to study the accumulation of americium in three species: the polychaete wormNereis diversicolor, the brackish-water amphipodGammarus duebeni and the harpacticoid copepodTisbe holothuriae. The species chosen are considered representative of groups having wide ecological importance. It was found that large differences in concentration factors occurred for the same organisms, depending upon aging of the contaminated medium; much higher and more variable values being found when uptake was from freshly contaminated solutions than from those aged up to a week. The interaction of specimens with physico-chemical reactions of americium which appear to take place within the first few days after its introduction into water are considered to be responsible for these differences. Uptake from contaminated water that had been allowed to age in the absence of organisms appears to be unaffected by subsequent conditioning by specimens. Americium concentration factors show a strong tendency to increase with decreasing size of the species, varying from over 1000 forT. holothuriae to about 3 forN. diversicolor. The possibility that the mechanisms regulating the uptake of actinides in different species may depend upon pH is briefly discussed.     

The hydrologic environment of Pool 19 of the Mississippi River

Invertebrate populations and biomass were studied over four years (1978–1981) in a coastal, saline lagoon in eastern England. Both temperature and salinities fluctuated sharply, with salinities as high as 72.4 gl⁻¹. Of 14 taxa recorded, only three occurred in significant numbers. Nereis diversicolor exhibited summer peaks over all four years, but there was a gradual trend of decline. Corophium volutator showed peaks in 1978 and 1979, but was largely absent thereafter. Chironomus salinarius showed a peak in numbers in 1978, but was rare in 1979 and largely absent in the following two years. The peak biomass was more than 4 000 mg m-2 in 1978, but was less than half this in subsequent years, when it consisted mainly of N. diversicolor. Laboratory experiments tended to confirm that high summer salinities were the cause of poor invertebrate success. The controlled ingress of brackish water into the lagoon from the adjacent estuary should result in a greater and more predictable production of invertebrates, which form the food supply of the chicks of an important colony of avocets Recurvirostra avocetta.     

Determination and localization of oil components in living benthic organisms by fluorescence microscopy

Microspectrofluorometric measurements are made to determine uptake and distribution of oil in marine organisms after exposure to crude oil. Equipment combining fluorescence microscopy with spectral analysis of the fluorescence emission is described. After contamination with oil, the intestine content ofLumbricillus lineatus, Nereis diversicolor andAnaitides mucosa shows a fluorescence emission at long wavelengths with a maximum at about 550 nm; this is in contrast to the fluorescence emission of these organisms without oil contamination. There is evidence that aromatic hydrocarbons are metabolized in the intestine of the worms studied.     

Phytoplankton succession affects the composition of Polynucleobacter subtypes in humic lakes

Phytoplankton influence the composition of bacterial communities, but the taxonomic specificity of algal–bacterial interactions is unclear due to the aggregation of ecologically distinct bacterial populations by community characterization methods. Here we examine whether phytoplankton seasonal succession affects the composition of subtypes within the cosmopolitan freshwater bacterial genus Polynucleobacter. Changes in the composition of Polynucleobacter subtypes were characterized in samples collected weekly from May to August in 2003 and 2008 from three humic lakes using terminal restriction fragment length polymorphism fingerprinting of the protein‐encoding cytochrome c oxidase ccoN gene. Changes in phytoplankton population abundances explained, on average, 30% of temporal variation in the composition of Polynucleobacter subtypes and the interaction between phytoplankton and the environment explained an additional 18% of temporal variation. The effect of phytoplankton on specific Polynucleobacter subtypes was experimentally confirmed by changes in Polynucleobacter subtype composition following incubation with different phytoplankton assemblages or a no‐phytoplankton control. Phytoplankton‐associated subtypes and differentiation in substrate use among subtypes likely contribute to the effects of phytoplankton on Polynucleobacter subtype composition. Interactions between unique Polynucleobacter populations and phytoplankton highlight the ecological significance and specificity of species interactions in freshwater communities.     

Development and validation of an experimental life support system for assessing the effects of global climate change and environmental contamination on estuarine and coastal marine benthic communities

An experimental life support system (ELSS) was constructed to study the interactive effects of multiple stressors on coastal and estuarine benthic communities, specifically perturbations driven by global climate change and anthropogenic environmental contamination. The ELSS allows researchers to control salinity, pH, temperature, ultraviolet radiation (UVR), tidal rhythms and exposure to selected contaminants. Unlike most microcosms previously described, our system enables true independent replication (including randomization). In addition to this, it can be assembled using commercially available materials and equipment, thereby facilitating the replication of identical experimental setups in different geographical locations. Here, we validate the reproducibility and environmental quality of the system by comparing chemical and biological parameters recorded in our ELSS with those prevalent in the natural environment. Water, sediment microbial community and ragworm (the polychaete Hediste diversicolor) samples were obtained from four microcosms after 57 days of operation. In general, average concentrations of dissolved inorganic nutrients (NO₃^?; NH₄⁺ and PO₄^?3) in the water column of the ELSS experimental control units were within the range of concentrations recorded in the natural environment. While some shifts in bacterial community composition were observed between in situ and ELSS sediment samples, the relative abundance of most metabolically active bacterial taxa appeared to be stable. In addition, ELSS operation did not significantly affect survival, oxidative stress and neurological biomarkers of the model organism Hediste diversicolor. The validation data indicate that this system can be used to assess independent or interactive effects of climate change and environmental contamination on benthic communities. Researchers will be able to simulate the effects of these stressors on processes driven by microbial communities, sediment and seawater chemistry and to evaluate potential consequences to sediment toxicity using model organisms such as Hediste diversicolor.     

Investigations of the anaerobic metabolism of the polychaete wormNereis diversicolor M.

Summary The anaerobic metabolism ofNereis diversicolor M. was studied during various periods of experimental anaerobiosis.The degradation of glycogen is shown to be the main source of anaerobic energy production. During first hours of anaerobiosis, aspartate, in addition to glycogen, is metabolized in considerable quantities.Five acids were found to accumulate as end-products: alanine, D-lactate, succinate, acetate and propionate (Table 2).Alanine is accumulated only during the first hours of anaerobiosis. The increase in alanine is correlated with a decrease in aspartate.D-Lactate is the main end-product during the first 24 h of anaerobiosis, and continues to be produced even during prolonged anaerobiosis. In accordance with lactate production,Nereis diversicolor possesses a high glycolytic capacity (Table 4).The major end-products of long term fermentation are propionate and acetate. In contrast to other end-products, these acids are excreted in substantial amounts.Abbreviations GAPDH glyceraldehydephosphate dehydrogenase, EC 1.2.1.12 - LDH lactate dehydrogenase, EC 1.1.1.27 - GOT aspartate aminotransferase, EC 2.6.1.1 - GPT alanine aminotransferase, EC 2.6.1.2 - MDH malate dehydrogenase, EC 1.1.1.37 Supported by Deutsche Forschungsgemeinschaft (Gr 456/5 and Gr 456/6)