Symbiotic bacteria in the nudibranch mollusk Dendrodoris nigra: fatty acid composition and ultrastructure analysis

In the present study, the hypothesis that marine nudibranch mollusks harbor symbiotic bacteria was tested using analyses of fatty acids as biochemical markers and transmission electron microscopy of the tissues of Dendrodoris nigra (Gastropoda/Opisthobranchia/Nudibranchia). An aberrant level of the odd-numbered carbon chain and branched fatty acids, iso- and anteiso- that are specific for bacteria, was detected in the nudibranch tissues. Their amounts in the notum exceeded significantly that in the viscera. Rod-shaped gram-negative bacteria were revealed in the epithelial cells of the notum and the mantle edge as well as in the adjoining glycocalix. These bacteria were enclosed in secondary vacuoles in the epithelial cells. The consequent stages of inoculation of the bacteria into the cytoplasm of epithelial cells, from adhesion to the apical surface to invagination of the cell membrane and formation of the vacuole with an enclosed bacterium, were observed. The presence of dividing bacteria suggests that the epithelium includes a renewable, dividing population of symbiotic bacteria. No bacteria were detected in the gonads and the digestive system. Probable functions of these symbiotic bacteria such as involvement in protection or defense from predators and environmental impacts as well as their nutritional role in the nudibranch are discussed.     

An evaluation of the ABC-method (abundance/biomass comparison) as applied to macrozoobenthic communities living on tidal flats in the Dutch Wadden Sea

The ABC-method proposed by Warwick (1986) for detecting pollution effects on marine zoobenthic communities, was tested on distribution patterns of numbers and biomass among species in macrozoobenthos samples taken annually for 13 yr in a uniform way at 15 tidal-flat stations in the western part of the Dutch Wadden Sea. Along the margins of the tidal-flat area studied, where exposition to either drainage or water movements is extreme, the k-dominance plots for numbers were generally situated above those for biomass. In these areas values for biomass and species richness were relatively low and a few small-sized species such asCorophium volutator andHydrobia ulvae were very numerous. In the central part of the area, where environmental conditions are less severe, values for biomass and species richness were higher. Biomass was dominated by large adults ofMya arenaria, Mytilus edulis and/orArenicola marina. However, in only a restricted part of this area were plots for numbers consistently below biomass plots. In most of the central area high numbers ofH. ulvae occasionally occurred, causing plots for numbers to be situated above those for biomass during such periods. There appears to be no reason to interprete a high abundance ofH. ulvae as a sign of pollution or any other kind of disturbance or stress. It is concluded that the ABC method cannot be applied to tidal flat communities without reference to long-term and spatial series of control samples. In areas where such smallsized and sometimes numerous species asH. ulvae occur in strongly fluctuating numbers, the method appears useless for assessing the pollution status of a benthic community.     

Inception of formation and early morphogenesis of the bacterial light organ of the sea urchin cardinalfish, <Emphasis Type="Italic">Siphamia versicolor</Emphasis>

The cardinalfish Siphamia versicolor (Perciformes: Apogonidae) forms a bioluminescent symbiosis with the marine luminous bacterium Photobacterium mandapamensis, harboring the bacteria in a ventral, disc-shaped light organ and using the bacterial light apparently for counterillumination and attracting prey. Little definitive information has been available on the developmental and microbiological events surrounding the initiation of symbiosis, a critical stage in the life history of the fish, in S. versicolor or any of the many other species of bacterially luminous fish. To identify the stage at which light organ formation begins, to determine the origin of cells forming the light organ, and to characterize its bacterial colonization status during development, early developmental stages of S. versicolor obtained and reared from wild-caught mouth-brooding males were examined with histological and microbiological methods. A light organ primordium was not evident in embryos, post-embryos, or pre-release larvae, whereas the light organ began to form within 1 day of release of full-term pre-flexion larvae from the mouths of male fish. Analysis of post-release larvae revealed that the light organ arises from a proliferation and differentiation of intestinal epithelial cells, and that it quickly develops structural complexity, including the formation of chambers and gaps contiguous with the intestinal epithelium. However, the nascent light organ remained uncolonized by the symbiotic bacteria through several days of post-release development, even in the presence of high numbers of the symbiotic bacteria. These results demonstrate that the inception of light organ formation in S. versicolor occurs independently of its symbiotic bacteria and that receptivity to bacterial colonization apparently requires substantial post-release development of the light organ. Larvae therefore most likely acquire their symbiotic bacteria from seawater, during or shortly after the transition from the pre-flexion to the flexion developmental stage.     

Gonadal development in the coral reef damselfish<Emphasis Type="Italic"> Dascyllus flavicaudus</Emphasis> from Moorea,French Polynesia

The present study investigated the sexual pattern of the yellow-tailed dascyllus, Dascyllus flavicaudus (Randall and Allen), through histological examination of the process of gonadal differentiation and maturation and through male-removal experiments in the laboratory. The study was conducted in Moorea, French Polynesia (17°32S; 149°50W), from 31 December 1994 to 11 March 1995. For gonadal histology, two populations were sampled, one consisting of large aggregations (n=54) and the other consisting of small- to medium-sized groups around isolated corals (n=55). An additional small sample (n=21) was also collected from a population that consisted of small groups around isolated corals. After an initially undifferentiated state, gonads of D. flavicaudus developed an ovarian lumen, followed by oocytes in the primary growth stage. From this ovarian state or from more developed ovaries with cortical-alveolus stage oocytes, some gonads developed into testes through degeneration of oocytes and development of spermatogenic tissue. In all three populations, spermatogenic tissue developed only in gonads that contained pre-vitellogenic oocytes (termed as mixed stage 1–2 gonads). The two main populations did not differ in their expression of sex change despite differences in their social organization. In both populations, size of individuals with mixed stage 1–2 gonads overlapped mainly with the size range of immature females, which suggests that functional female-to-male sex change was rare. The hypothesis that D. flavicaudus is primarily gonochoristic is further supported by removal experiments in laboratory groups, in which removal of a dominant male(s) failed to induce sex change in any of the high-ranking females. Yet, in all three field populations, some fish with mixed stage 1–2 gonads were found within the size ranges of mature females, which indicates possible occurrences of functional sex change. These conflicting results indicate that it may be premature to draw any definitive conclusions about the sexual pattern of D. flavicaudus.Communicated by T. Ikeda, Hakodate     

Anatomical and experimental evidence for particulate feeding in<Emphasis Type="Italic"> Lucinoma aequizonata</Emphasis> and<Emphasis Type="Italic"> Parvilucina tenuisculpta</Emphasis> (Bivalvia: Lucinidae) from the Santa Barbara Basin

Previous nutritional models for adults of the lucinid bivalve Lucinoma aequizonata contend that symbiotic chemoautotrophic bacteria provide most of the organic carbon for the host. The existence of this symbiosis, coupled with the hosts distinctive anatomical features, shaped the impression that particulate feeding was not a significant part of L. aequizonata nutrition. Here, we use several techniques to show that particulate feeding is a consistent and important part of the L. aequizonata nutritional strategy. Histological and scanning electron microscopy observations reveal that the gills of L. aequizonata, like those of the lucinid Parvilucina tenuisculpta, have functional mucociliary epithelia, able to transport captured particles to the mouth. Observations of gut content and radiolabeled feeding experiments indicate that L. aequizonata does ingest and assimilate carbon from particulate organic matter. Furthermore, molecular identification of a broad spectrum of organisms in the guts of native adult specimens demonstrates that L. aequizonata is non-selective when ingesting organic material, and has a mixotrophic diet.Electronic Supplementary Material Supplementary material is available in the online version of this article at Communicated by P.W. Sammarco, Chauvin     

Long-term changes in reproductive patterns of the holothurian<Emphasis Type="Italic"> Oneirophanta mutabilis</Emphasis> from the Porcupine Abyssal Plain

Time-series studies (1989–2002) in the NE Atlantic have shown large-scale changes in the composition and structure of the benthic community on the Porcupine Abyssal Plain. Radical changes in the abundance of some species in 1996 led to a significant shift in the way in which organic matter was reprocessed at the seabed. This article examines the reproductive processes of the holothurian Oneirophanta mutabilis collected during the time series. The reproductive biology of O. mutabilis is reviewed. No males were evident in any of the samples. The sex ratio (females: no gender) was biased significantly towards no-gender individuals. The maximum egg size was 650 m and there was no evidence of synchrony in reproduction. Significant changes in the oocyte-size distribution and the fecundity of O. mutabilis were noted through time, coinciding with the time of greatest faunal change in the benthic community. There was an increase in the proportion of previtellogenic oocytes and a decrease in the proportion of mature vitellogenic oocytes in 1997 and 1998, in parallel with a significant decrease in fecundity of the post-1996 samples. Samples from 2002 showed a reversal in the reproductive trends, with an increase in the proportion of mature vitellogenic oocytes and fecundity. The results are discussed in relation to large changes in abundance of the epibenthos on the Porcupine Abyssal Plain. It is suggested that the superabundance of certain megafaunal species on the abyssal seafloor affected the availability of trophic resources for O. mutabilis, leading to the changes in its reproductive effort.     

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.     

Sexual reproduction of the photosymbiotic ascidian <Emphasis Type="Italic">Diplosoma virens</Emphasis> in the Ryukyu Archipelago,Japan: vertical transmission,seasonal change,and possible impact of parasitic copepods

Diplosoma virens is a colonial ascidian hosting prokaryotic algae Prochloron sp. in the common cloacal cavity of the colonies and is sometimes parasitized by notodelphyid copepods. In ascidian–Prochloron symbiosis, it is generally known that the host larvae acquire the algal symbionts from their mother colonies to maintain the symbiosis. A histological study of the sexually mature colonies of D. virens showed that the algal symbionts attach to pre-hatching larvae on the rastrum (plant rake) projected from the postero-dorsal part of the larval trunk, and then the rastrum is packed in the posterior half of the larval trunk that will become a cloacal cavity after metamorphosis. This process is the same as that of D. simile. Monthly sampling of D. virens colonies showed that they have embryos in summer in Ryukyus, situated near the northern-most limit of the coral reefs in the West Pacific. While the frequencies of copepod parasitism were variable among the populations, the colonies from a highly parasitized population had a significantly smaller number of eggs/embryos per zooid than the colonies from the less parasitized populations. The parasites probably have an inhibitory impact on the sexual reproduction of the host colonies.Communicated by T. Ikeda, Hakodate     

Symbiosis within a symbiosis: Intracellular bacteria within the endosymbiotic protistNephromyces

The marine protistNephromyces Giard, 1888 is a chronic endosymbiont of molgulid tunicates. Ultrastructural and cytochemical studies of this protist, isolated from molgulid hosts collected from the Pacific, Atlantic and Gulf Coasts of the United States, indicate thatNephromyces is itself chronically infected with Gramnegative, intracellular bacteria. Molgulid tunicates are thus the locus of a nested,tripartite endosymbiosis. Intracellular bacteria are present in both trophic and reproductive stages ofNephromyces, suggesting that the bacterial-Nephromyces symbiosis is an hereditary association. The presence of endosymbionts inNephromyces raises the possibility that some ofNephromyces' metabolic characteristics, in particular its high urate oxidase activity, might be supplied not byNephromyces itself, but rather by its intracellular bacteria, possibly functioning as peroxisomal analogues.     

Sulfide-oxidizing bacteria in the burrowing echinoid,Echinocardium cordatum (Echinodermata)

Symbiotic filamentous bacteria thrive in the intestinal caecum of the deposit-feeding echinoid Echinocardium cordatum. Specimens of E. cordatum were collected at Wimereux (Nord Pas-de-Calais, France) in 1991. Their symbiotic bacteria build nodules by forming multilayered mats around detrital particles that enter the caecum. The morphological features of the bacteria are those of Thiothrix, a sulfide-oxidizing genus. The filaments, which may form rosettes, are sheathed and made by a succession of hundreds of rod-shaped bacteria which store elemental sulfur in the presence of external sulfide. Live bacteria are restricted to the outer layers of the nodules. Their sulfide-oxidizing activity was investigated, using a Biological Oxygen Monitor, by measuring the O₂-consumption when reduced sulfur compounds are provided. They oxidize thiosulfate and sulfide. Optimal sulfide oxidation occurs at intermediary pO₂ (100 to 160 M O₂l^-1). Spectrophotometry has confirmed that the sulfur content of the filamentous symbiotic sulfideoxidizing bacteria depends on the presence of external sulfide. This is the first report of symbiotic intradigestive Thiothrixspp.-like bacteria; it lengthens the list of symbioses between sulfide-oxidizing bacteria and invertebrates from sulfide-rich habitats.     

Larval metamorphosis of the sea urchins,<Emphasis Type="Italic"> Pseudocentrotus depressus</Emphasis> and<Emphasis Type="Italic"> Anthocidaris crassispina</Emphasis> in response to microbial films

In Japan, mass-production of sea urchin juveniles involves the culture of periphytic diatom films on plastic plates in 5- to 15-tonne tanks for the induction of larval metamorphosis. This study focused on the larval response of sea urchins, Pseudocentrotus depressus and Anthocidaris crassispina, to natural microbial films in the sea and diatom-based films formed in the tanks. The effect of diatoms and bacteria on larval metamorphosis was also examined using laboratory-cultured diatom-based films in the presence of germanium dioxide and antibiotics during film culture. Furthermore, the nature of the cue of the cultured diatom-based film was also investigated. Results showed that P. depressus and A. crassispina metamorphosed both on natural microbial films and diatom-based films in a tank. In the sea, the metamorphosis (%) of P. depressus increased gradually in accordance with the immersion period of film formed on glass slides, whereas the larval metamorphosis of A. crassispina had a bell-shaped response curve. In the tank, although the diatom-based films showed a low inducing activity for larval metamorphosis of A. crassispina, the metamorphosis of P. depressus larvae increased linearly in accordance with the diatom density. These results suggest that diatom-based films could promote the larval metamorphosis of P. depressus, but are less important in A. crassispina. In a simultaneous larval assay (May), P. depressus showed a higher percentage of metamorphosis than A. crassispina. We concluded that the former is more sensitive to diatom-based film than the latter and that this is due to differences in their natural habitats. For laboratory-cultured diatom-based film, both species of sea urchins showed a similar response, in which reduction in diatom and bacteria density resulted in a decrease in the original inducing activity. There seems to be a synergistic effect between diatom and bacteria in inducing larval metamorphosis. Films subjected to treatment with 0.1 N HCl were no longer inductive for either sea urchin, while those films treated with 40^°C heat or EtOH (5% and 10% EtOH) showed a significant reduction in the inducing activity. Thus the surface-associated cues may be highly susceptible to the above treatments.Communicated by T. Ikeda, Hakodate     

The reproductive biology of Hymenaster membranaceus from the Rockall Trough,North-East Atlantic Ocean,with notes on H. gennaeus

Samples of the deep-sea spinulosan asteroid Hymenaster membranaceus Wyville Thomson were collected in a timeseries of 19 bottom trawls spanning the period April 1978 to October 1981 from a 2200 m-deep station in the northern Rockall Trough. The reproductive biology of this species was studied from histological sections of the gonad, and compared with that of H. gennaeus H. L. Clark which was collected in the same hauls. At first sexual development, oogonia develop in nests surrounded by small accessory cells. Previtellogenic oocytes remain in the periphery but, at maturity, oocytes ranging up to 1 100 m fill the ovary. A variety of accessory cells pack the lumen and may be nutritive or degenerative. It appears that a small number of oocytes are spawned intermittently, but there is no evidence of overproduction and break-down of superfluous oocytes. A few large oocytes become senescent and undergo internal break-down, releasing periodic acid Schiff-positive material into the lumen. Size-frequencies of oocytes indicate that eggs may be spawned as a continuous slow release, and there is no evidence of reproductive synchrony between or within samples. On reaching maturity, males appear always to be ready to release spermatozoa. Spawning is probably stimulated by egg release during chance encounters with mature females. There is no evidence for brooding, and from the large size and yolky nature of the egg direct lecithotrophic development at or near the seabed is inferred. A limited histological study of H. gennaeus indicates that egg production is very similar, but the two species differ in the nature of the accessory cells and amorphous material filling the lumen.     

Spawning and development in <Emphasis Type="Italic">Osedax</Emphasis> boneworms (Siboglinidae,Annelida)

We report observations on spawning and early development in bone-eating worms of the genus Osedax. Individual females of Osedax rubiplumus were observed at 1820 m depth freely spawning hundreds of oocytes, and females of an undescribed species, Osedax “orange collar”, were observed spawning in laboratory aquaria. Cytological and molecular analysis of the spawned oocytes of two Osedax species revealed no evidence for the bacterial endosymbionts that the female worms require for their nutrition, suggesting that the bacteria must be acquired later from the environment, as they are in other siboglinids. Individual O. “orange collar” females released an average of 335 (±130) eggs per day, but the number of oocytes spawned per day varied greatly, suggesting that not all the females spawned daily. Fertilization rates of the spawned oocytes varied from 0 to 100%, though most females showed nearly 100% fertilization rates. Oocytes spawned in the laboratory at 4–6°C were negatively buoyant. If fertilized, these oocytes extruded polar bodies and then after at least four hours cleaved unequally. Subsequent cleavages occurred in a spiral pattern at roughly 2-h intervals, resulting in free-swimming trochophore larvae after 24 h. These lecithotrophic trochophores swam for 9–16 days before settling with several hooked chaetae, similar to those of dwarf Osedax males. The larval life span of the Osedax species studied in the laboratory appears to be shorter than in closely related Vestimentifera. Osedax rubiplumus, on the other hand, has much larger oocytes and so may have greater dispersal potential than these other Osedax species. The high fecundity and apparently continuous reproduction of Osedax boneworms permits the opportunistic exploitation of sunken vertebrate bones. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Carrion-feeding on the sediment surface at nocturnal low tides by the polychaete<Emphasis Type="Italic"> Phyllodoce mucosa</Emphasis>

Harsh physical conditions in the intertidal zone are the cause of an ample amount of dead macroinvertebrates, which constitute a food source for carrion-feeders. In the European Wadden Sea, this trophic guild includes decapod crustaceans and fish when the tide is in, while during nocturnal low tides the polychaete Phyllodoce mucosa is attracted in large numbers by dead mollusks, crabs or worms on the sediment surface. Within 10 s worms emerged to the surface, crawled as far as 15 m on mucus trails towards the carcass, sucked in tissue up to one-third of their own weight, and then quickly retreated to below the surface. Abundance of P. mucosa was highest in the lower intertidal zone and winter. The seaward high abundance pattern, however, did not continue into the shallow subtidal. In summer, few were attracted during daytime or when the tide was in. However, up to 447 worms aggregated at a single crushed mussel within 20 min at dusk during low-tide exposure. This study suggests that during winter carrion-feeding is an important trophic niche on cold-temperate, intertidal mud flats occupied by a phyllodocid polychaete that is segregated in feeding time from most other scavengers and benefits from cold-sensitive benthic invertebrates.Communicated by O. Kinne, Oldendorf/Luhe     

The trophic link between squid and the emperor penguin <Emphasis Type="Italic">Aptenodytes forsteri</Emphasis> at Pointe Géologie,Antarctica

Cephalopod beaks retrieved from stomachs of dead emperor penguin chicks at Pointe Géologie, Terre Adélie, provide information on taxonomic and size composition of the penguin’s squid diet, on the trophic range of the squid species preyed upon and on the fractional trophic impact of the penguin on the whole food web. Emperor penguins prey upon four squid species (Psychroteuthis glacialis, Kondakovia longimana, Gonatus antarcticus, Alluroteuthis antarcticus) and do not take squid larger than 480 mm mantle length. Larger squid live either below the penguin’s diving range or are beyond its handling capacity. Nitrogen stable isotope ratios indicate that squids cover a range of about two trophic levels (2.5–8‰ δ¹⁵N). The impact of the emperor penguin, however, concentrates on the upper part of this range, about 68% of its squid prey being >6‰ δ¹⁵N. The principal components of the emperor’s diet, fish, krill and squid, differ distinctly in average trophic level. Consequently the trophic position of the emperor penguin changes accordingly with diet composition and may differ by almost one trophic level between different emperor penguin colonies.     

Ultrastructure of post-fertilization development in the red alga<Emphasis Type="Italic"> Callophyllis linearis</Emphasis> (Kallymeniaceae,Cryptonemiales, Rhodophyta)

The ultrastructure of post-fertilization development of the red alga Callophyllis linearis (Kyl.) Abb. & Norr. of the Kallymeniaceae showed that the Kallymeniaceae are more closely related to the Cryptomeniales than to the Gigartinales. The auxiliary cell produces multinucleate gonimoblast initials which divide to produce generative gonimoblast cells. These cells undergo repeated cleavages to form clusters of carpospore initials, which finally differentiate into carpospores. Carpospore differentiation is simple and occurs in three developmental stages. In the first stage, young carpospores are avacuolate, contain a large nucleus, have few undeveloped plastids and exhibit incipient dictyosome activity. During the intermediate stage, plastids develop their internal thylakoid system, starch granules are formed and dictyosomes continue to produce vesicles that contribute carpospore wall material. The formation of fibrous vacuoles originates from the fibrous vacuole associated organelles (FVAOs). In the final stage, mature carpospores form cored vesicles. They also contain well-developed plastids, fibrous vacuoles, numerous starch granules and are surrounded by a two-layered carpospore wall.Communicated by O. Kinne, Oldendorf/Luhe     

Reproductive strategies of two deep-sea gastropod species from the Porcupine Seabight (Northeast Atlantic)

Two species of small gastropods (<6 mm in length), Amphissa acutecostata (Philippi, 1844) and Gymnobela subaraneosa (Dautzenberg and Fischer, 1896), widely distributed in the northeast Atlantic, were found in large numbers in the Porcupine Seabight (Northeast Atlantic). Except for some aspects of taxonomy and distribution, as well as some data on larval development, the biology of these species is unknown. This study describes basic aspects of the life-history strategies of both species. Histological studies showed that oocyte and sperm development in both species was similar to the gametogenetic patterns observed in other deep-sea gastropods. In females, oogonia proliferated in the germinal epithelium and developed into previtellogenic oocytes (30–40 m), which grew into vitellogenic primary oocytes. Vitellogenic oocytes were covered by a thin layer of follicle cells involved in the vitellogenic processes. The maximum size for mature oocytes was 99.06 m for A. acutecostata and 114.82 m for G. subaraneosa. In A. acutecostata most of the volume of the ovary was occupied by previtellogenic and early vitellogenic oocytes, whereas in G. subaraneosa most of the volume was filled by large vitellogenic oocytes. Both species showed quasi-continuous production of oocytes. The oocyte size-frequency diagrams suggested a continuous release of a small number of oocytes throughout the year for A. acutecostata, and asynchronous periodic spawning events for G. subaraneosa. Gonad development and gametogenesis could be strongly affected by presence of parasites in one of the populations of G. subaraneosa.Communicated by J.P. Thorpe, Port Erin     

Gametogenesis, spawning behavior, and early development in the “iceworm”Hesiocaeca methanicola (Polychaeta: Hesionidae) from methane hydrates in the Gulf of Mexico

Gametogenesis, spawning behavior, and early development have been described in the methane-seep polychaete Hesiocaecamethanicola Desbruyères and Toulmond, 1998, the first documentation of the reproductive biology of any cold-seep polychaete species. The worms were collected at the Green Canyon site in the Gulf of Mexico (27°44.77N; 91°13.33W) in August 1997. The gonads are situated in the neuropodia in association with an intraepithelial capillary system. Oogenesis is intraovarian, with oocytes remaining in the ovary until late vitellogenesis when they are released into the coelom for final growth and differentiation. Vitellogenesis involves both autosynthetic and heterosynthetic processes and, along with ovarian structure, suggests a relatively slow process of egg formation. Sperm differentiation is intertesticular until early spermiogenesis, when spermatid tetrads complete development in the coelom. The mature spermatozoon is an ect-aquasperm consisting of a tapering, digitate acrosome, a spherical nucleus, a midpiece containing from six to eight mitochondria and glycogen stores, and two centrioles associated with the flagellum. Both sexes undergo broadcast spawning, but the males spawn exclusively through the anus, a behavior previously unknown in polychaetes. Artificial fertilization resulted in spiral cleavage and development to the trochophore stage. Oocyte size-frequency analysis of adult females indicates a wide range of oocyte sizes and vitellogenic stages, suggesting that asynchronous gametogenesis occurs. Received: 13 July 2000 / Accepted: 29 November 2000     

Bacterial symbiosis in Northeast Pacific Vestimentifera: a TEM study

A number of new vestimentiferan species occur at northeast Pacific hydrothermal vent sites. The trophosome and bacterial symbionts of three species, collected from the Juan de Fuca and Explorer Ridges between 1984 and 1986, were studied by transmission electron microscopy (TEM). As in Riftia pachyptila, trophosome tissue is organised into lobules each having an axial blood vessel, and intracellular bacterial symbionts are contained in membrane vacuoles. The bacteria have many cytoplasmic inclusions including tubular membrane systems, glycogen-like particles and poly--hydroxybutyrate (PHB) or sulfur bodies. Glycogen production may be quantitatively important to both the symbionts and the host. Glycogen-like granules appear to first accumulate in the bacterial cells and then be released into the bacteriocyte cytoplasm as bacteria are degraded. Although various stages of bacterial growth and degradation are observed, data are insufficient to verify any across-lobule progression of these processes. Morphological comparison of the symbionts reveals that similar symbionts are found in different vestimentiferan species and that one to two bacterial types exist within single individuals. Two possible models of trophosome function and nutrient exchange are discussed.Deceased     

POM in macro-/meiofaunal food webs associated with three flow regimes at deep-sea hydrothermal vents on Axial Volcano,Juan de Fuca Ridge

Deep-sea hydrothermal vent ecosystems host both symbiotic and non-symbiotic invertebrates. The non-symbiotic vent fauna is generally assumed to rely on free-living chemoautotrophic bacteria as their main food source but other sources such as detritus have recently been suggested to be a part of the invertebrate diets. Little is known about how food availability influences the distribution of vent organisms on a small scale. In addition, the feeding ecology and role of small, often numerically dominant invertebrates, the meiofauna is poorly understood at vents. In this study, we used stable carbon and nitrogen isotopic analysis to investigate the role of particulate detritus in the diets of macro- and meiobenthic invertebrates within three vent assemblages at Axial Volcano, Juan de Fuca Ridge, and Northeast Pacific. Particulate organic matter of a detrital origin became more important in the diet of invertebrates in assemblages typically associated with low-hydrothermal flow intensities. Meiobenthic species occupied several different feeding guilds and trophic levels in the assemblages investigated. We conclude that small-scale spatial variability in food sources is an important feature of vent food webs and that spatial patterns observed here and elsewhere are shaped by variations in hydrothermal discharge.