Structure and function of the third maxillipeds of the banana prawn Penaeus merguiensis

The functions of the third maxilliped of Penaeus merguiensis de Man are described. It has a 6-segmented endopodite which is used in feeding, grooming and possible pheromone reception. In feeding, large pieces of food are gripped by the stout spines on inner borders of the ischia which hold them to the mouth so that particles can be torn off and ingested. All the distal segments have long setae used in grooming chelate pereiopods and the antennules. In mature males there is a tuft of about 300 slender setae at the end of the propus, but its function remains obscure.     

Plasticity of biometrical and mechanical properties of Echinocardium cordatum spines according to environment

We investigated the plasticity of spines of the burrowing heart urchin Echinocardium cordatum inhabiting sandy beaches. This urchin has very specialized spines which play specific roles in feeding and locomotion. Biometrical and mechanical properties of the spine skeleton of individuals from a wave-exposed intertidal site and a protected subtidal site (Brittany, France) were compared and related to sediment grain size, food supply and urchin small-scale distribution and burrowing depth. The intertidal site was characterized by coarser sand; the urchins were more scattered and more deeply buried than in the subtidal site. Intertidal urchins were bigger than subtidal urchins but showed similar resource allocation. Spines of intertidal urchins were longer than those of subtidal urchins, their architecture was reinforced and they presented a higher fracture force and stiffness. These spine differences would be essentially due to passive plasticity, but could also be adaptative at a certain extent.     

Growth of the deep-sea irregular sea urchins Echinosigra phiale and Hemiaster expergitus in the Rockall Trough (N.E. Atlantic Ocean)

Growth in the deep-sea irregular sea urchins Echinosigra phiale (family Pourtalesiidae) and Hemiaster expergitus (family Hemiasteridae) was studied from deep-sea samples taken during the years 1973 to 1985 from two stations at 2900 and 2200 m depth in the Rockall Trough (N.E. Atlantic Ocean). Growth zones, similar to those described from sea urchins in shallow water, are present as a series of wide white bands separated by narrow, dark rings in the calcite stereom of the test plates after heating to 350°C. In shallow water, such growth zones seem to result from seasonally varying growth rates. In the supposedly constant conditions in the deep sea, a seasonal growth pattern is unexpected but may occur in response to recently discovered annual pulses in downward flux of detritus from the euphotic zone, providing a seasonally varying food supply for such deposit-feeding species living in the bottom sediment. On this assumption, growth curves were fitted to counts of growth zones (as representing age in years), in the larger lateral and ventral test plates of E. phiale and H. expergitus. The opportunity was also taken to fit growth curves derived from counts of growth zones in samples of the inshore spatangoids Spatangus purpureus and Echinocardium pennatifidum. Plots of counts against test length of Echinosigra phiale and H. expergitus, although scattered and not clearly asymptotic, indicate, growth to be slower than in the two inshore spatangoids, and than in the coastal species Echinocardium cordatum, for which there are good recent growth data, available.     

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.     

Functional morphology and feeding behavior of Paraprionospio pinnata (Polychaeta: Spionidae)

The functional morphology and feeding behavior of Paraprionospio pinnata (Ehlers) were studied. The tentacular feeding palps of P. pinnata possess four groups of cilia: laterals, latero-frontal cirri, frontals and basal transverse rows. The lateral cilia beat in metachronal waves creating in current that flows toward the frontal surface of the palp. The latero-frontal cirri deflect suspended particles onto the frontal surface and potentially resuspend deposited particles. The frontal cilia line the groove of the palp and transport particles to the ciliated pharynx. The basal transverse cilia also beat in metachronal waves and together with the frontal cilia of the branchiae produce a U-shaped current that removes particles rejected by the pharynx from the burrow. The significance of particle selection at the site of the pharynx in spionid polychaetes is hypothesized to be constrained by the interaction of specific morphologies and behaviors that reduce the effects of epifaunal predation and browsing. The relationship between sediment permeability and type of respiratory or ventilation current is hypothesized to be a potentially important factor in understanding the distribution of spionid polychaetes.     

The effect of food supply on feeding strategy in sessile female gastropods<Emphasis Type="Italic"> Crepidula fecunda</Emphasis>

Sessile females of the gastropod Crepidula fecunda are exclusively suspension-feeders. Our previous work showed that particles removed from suspension are entrained on the gill and continuously accumulate as a mucous string on the distal margin of the lamella, although some are transferred by a ciliary tract to the food pouch, where they form a mucous ball. The mucous string is transferred from the gill margin to the neck canal, where it forms a mucous cord which is transported to the mouth. The mucous ball and the mucous cord are ingested with the aid of the radula. In this study we examined the response of the system to a wide range of cell concentrations of the flagellate Isochrysis galbana. As the particle concentration increased from 5,000 to 200,000 cells ml^-1, the volume of the mucous ball almost doubled, as did the number of radular extrusions required to ingest it. The total volume of mucous balls produced and ingested increased very slightly as particle concentration rose from 5,000 to 100,000 cells ml^-1, but the volume of mucous cord material ingested increased four to five times over the same range. At higher food concentrations the mucous cord increased in thickness, but not in length, which is constrained by the length of the gill margin. At all particle concentrations the mucous cord accounted for 80–95% of the total material (mucous cords plus mucous balls). Ingestion rate reached a maximum at 140,000 cells ml^-1 for mucous balls and mucous cords. At higher concentrations the production of pseudofaeces increased from all sources ("type-I" pseudofaeces from rejected mucous balls; "type-II" pseudofaeces from rejected fragments of mucous cords; and "type-III" pseudofaeces, which come from particles trapped in a coarse mucous net at the mouth of the inhalant mantle cavity and are transferred via the lateral ciliary tract to the mantle margin, by-passing the food pouch). Thus C. fecunda has adopted a feeding strategy in which particle entrainment by the gills is continuous, even at very high ambient particle concentrations (200,000 cells ml^-1 for I. galbana), and excess material which cannot be ingested is rejected by one or more of three routes for the elimination of pseudofaeces.Communicated by P.W. Sammarco, Chauvin     

Bilaterality in the regular sea urchin Anthocidaris crassispina is related to efficient defense not to efficient locomotion

Regular sea urchins show pentaradial symmetry: they have round bodies made of five almost identical sectors. There is no preference in which sector should be anterior in their locomotion. Anthocidaris crassispina is exceptional among regular sea urchins. Morphometry showed that their tests had radial symmetry but the overall body shape showed bilateral symmetry because of the uneven distribution of spine length. This bilateral axis did not correspond to the ambulacrum III—interambulacrum 5 axis, which is the bilateral axis of irregular sea urchins. Their behavior in relation to this bilaterality was observed in the aquarium. At rest, most of them stayed in the upper corner of the aquarium keeping the sector with the shortest spine length (the shortest sector) toward the water surface. At walking, most of them moved on the vertical wall just under the water surface keeping the shortest sector directing to the water surface. When they walked in the open space, they moved in a random direction with respect to the bilaterality of the spines. The bilaterality and the behavior of this sea urchin were interpreted as the adaptation to defense not to efficient locomotion.     

Tube-building by a marine meiobenthic harpacticoid copepod

Pseudostenhelia wellsi Coull and Fleeger is a meiobenthic harpacticoid copepod inhabiting muddy, estuarine sediments. All individuals observed, and all ages including nauplii, build and inhabit elongate, mucous tubes which may extend to a depth of 3.9 mm into the sediment. Many of the narrow tubes, 0.27 mm in diameter, have a tube-cap extending 0.32 mm above the sediment-water interface. The tubes are a matrix of fine silt, sand and detritus cemented with a mucopolysaccharide, as shown by the Periodic Acid Schiff stain, apparently secreted from glands in the ventrolateral margin of the cephalothorax. Upon addition to sterile, homogeneous sediment, P. wellsi quickly (within 1 to 2 h) transform the upper 0.4 cm to a cohesive conglomerate of tubes, silt-clay particles and mucous filaments.     

Particularités microstructurales du squelette de Paracentrotus lividus et Arbacia lixula: Rapports avec l'écologie et l'éthologie de ces échinoïdes

Scanning-electron-microscope investigations on the test microstructure of two regular sea urchins, Paracentrotus lividus (Lamarck) and Arbacia lixula Linné have been especially devoted to the sutures between test plates, and the spines. Some features of both these components may be related to ecological and ethological differences between the two species. It is well documented that the structure of the sea urchin's test plates consists of a meshwork of calcareous trabecules embedded within the mesenchyme. The suture between the two lines of plates of each radial or intertradial zone exhibits a gap which plays some part in the process of the plates' growth. However, the gap may also constitute a complementary stress-breaker of mechanical forces (such as waves) which are exerted upon the test in the natural environment. In A. lixula this gap is so wide and the height: diameter ratio so low, that this species is particularly well fitted to bear physical stress and force on its apical region: since A. lixula almost exclusively inhabits vertical or subvertical rocky substrates at 2 to 15 m depth, where wave action is mainly exerted perpendicular to the substrate, these particular features of sutures and test shape may be a morphofunctional adaptation to this habitat. The sutures of P. lividus exhibit a narrower gap, making this species less able to bear strong apical pressure; consequently, P. lividus usually occurs on exposed horizontal or gently inclined substrates, but also inhabits sea-grass beds as deep as 15 to 20 m. The spines of P. lividus bear deep longitudinal grooves with lateral teeth, which seem especially fitted for collecting and transporting organic particles from the top of the spines to the apical region of the test where they are digested and assimilated by coelomocytes and epithelial cells. In the natural environment, the slightest water motion provides the spines with suspended particles; in extremely sheltered places or in aquaria however, the spines actively collect particles. Therefore, P. lividus populations can thrive in places where food resources other than suspended particles are scarce. The grooves on the spines of A. lixula are less marked, and thus unsuitable for collection of suspended particles. This species can therefore fulfil its energy requirements only by grazing and absorption of dissolved material. It appears that although belonging to the same biocoenosis, P. lividus and A. lixula do not occupy the same ecological niche. Their potential utilization in either urban or chemical pollution monitoring studies is discussed.     

Hemolysins from the mucus of Spirographis spallanzani (Polychaeta: Sabellidae)

Specimens of Spirographis spallanzani (Polychaeta: Sabellidae) were collected in the Gulf of Taranto, Italy, in 1991, and their mucous secretion was examined. The mucus possesses thermolabile biologically active proteins which produce hemolysis in vertebrate erythrocytes. The molecules are ineffective against both unfertilized and fertilized echinoid eggs and spermatozoa. The lytic reaction against rabbit erythrocytes varies as a function of cations, pH and temperature, and displays a fast time-course. No immunological cross-reaction could be demonstrated between hemolysins of S. spallanzani and Eisenia foetida andrei. SDS-PAGE analysis of mucus proteins reacting with sheep erythrocytes revealed that at least three major (16, 22 and 40 kDa) and two minor (29 and 34 kDa) components interact with membrane targets.     

Functional morphology of the podia and ambulacral grooves of the comatulid crinoid Antedon bifida (Echinodermata)

Feeding units — viz. triplets of unequally-sized podia associated with protective lappets — occur all along the pinnules of adult Antedon bifida (Pennant). Small food particles are trapped by direct mucus impingement to the wall of primary and secondary podia (there are no mucus net or mucus thread helping in this process). Large particles are caught by primary and secondary podia which partly curl over them. Small particles accumulate on the collecting podia before being transferred to the groove, while large particles are transferred one by one. Transfer of particles to the groove occurs by wiping the collecting podia on the ciliary tracts against the ciliary current. When active, tertiary podia always paddle against the ciliary current. They serve in bolus formation (mucus embedding of food particles), but do not participate in bolus compaction or propulsion. Elimination of unwanted particles occur through the action of secondary podia whose movements may disrupt the lappet's palisade, thus creating a sideward current that moves particles from the groove to the outside. Typical podial triplets do not occur along brachial and calycinal grooves. Both brachial and calycinal podia function mostly in guiding and regulating particle flow. The feeding structures of early pentacrinoid larvae of Antedon bifida recall to mind those of pinnules of adult individuals. They consist of twenty-five podia arranged in five radial triplets alternating with five interradial pairs. They are similar to the pinnular feeding structures of adults in that they have both collecting (radial) and paddling (interradial) podia.     

Feeding of the copepod Acartia tonsa on the diatom Nitzschia closterium and brown algae (Fucus vesiculosus) detritus

Integumental sensilla were examined in 39 species of meso- and bathypelagic shrimps and 6 species of epibenthic shrimps. A tuft organ, consisting of a group of openended, tubular setae and so probably functioning as a chemosensor, is described. A single tuft with ancillary smaller tufts occur on the dorsal median surface of the fourth abdominal segment and two pairs of tufts occur on the proximal dorsal region of the telson of all oplophorid and pandalid shrimps examined. An additional tuft is present on the fifth abdominal segment of the rare bathypelagic Physetocaris microphthalma Chace. No tuft organs were identified in Processa canaliculata Leach, suggesting that they may not occur in all families of caridean shrimps. Acanthephyrid, systellaspid and pandalid shrimps have the integument completely covered by scales. These are extremely delicate and are normally removed during the process of catching the shrimp through abrasion of the integument by the net. The scales are peculiarly orientated, pointing anteriorly in the anterior half and posteriorly in the posterior half of the body. They probably have a sensory function as distance receptors, monitoring water currents and disturbances in the environment surrounding the shrimp.     

Ammonia as confounding factor in toxicity tests with the sea urchin Paracentrotus lividus (Lmk)

The aim of this study was to investigate the toxicity of total ammonia towards sea urchin bioassays, in order to elucidate the role of ammonia as confounding factor in sediment quality assessment studies. New toxicity data, expressed as EC50, NOEC and LOEC are reported for Paracentrotus lividus based on experiments at differing pH values. Results confirmed that sperm cell toxicity test is not very sensitive to ammonia, while total ammonia embryotoxicity is strictly pH dependent. Accurate observations of embryotoxic effects at increasing total ammonia concentrations evidenced progressive shifts from malformed plutei to gastrula and blastula blockages. Toxicity data obtained for P. lividus are comparable with previous results with other echinoid species.     

Mode of particle ingestion in five species of suspension-feeding bivalve molluscs

In order to elucidate the mode of particle ingestion and the functional anatomy of the oesophagus in bivalves, a histological study was performed onMytilus edulis (Mytilidae),Crassostrea virginica (Ostreidae),Placopecten magellanicus, Chlamys varia, and juvenilePecten maximus (Pectinidae). Specimens were sampled from various sites in New Brunswick, Canada, and Brittany, France, from 1987 to 1989. The buccal, peribuccal, and oesophageal epithelia of all species contained a dense distribution of actively secreting mucocytes, although these were somewhat less abundant inCrassostrea virginica, which also has the shortest oesophagus. Mucocyte morphology, while constant within a family (Pectinidae), showed clear differences among families. Both acid and neutral mucopolysaccharides were secreted by the epithelial mucocytes of all species. Mucus and mucus-particle masses were observed in the peribuccal and buccal regions, as well as in the oesophageal lumina of all species, even in those specimens which had been maintained without feeding (Placopecten magellanicus) or held out of water for 48 h (C. virginica) prior to dissection and fixation. These results indicate that a basal level of mucus production and transport is continuous on the peribuccal, buccal, and oesophageal ciliated epithelia, regardless of the particle concentration in the external medium. Buccooesophageal glands, generally thought to be absent in the Bivalvia, were observed in one of the species examined (M. edulis). It is concluded that the mode of particle ingestion in these suspension-feeding bivalves is via ciliatransported mucus masses; the presence of buccooesophageal glands inM. edulis suggests a digestive role for the oesophagus in this species.     

Penetration and storage of sponge spicules in tissues and coelom of spongivorous echinoids

When echinoids feed on sponges, silicate spicules of the sponge were found to enter their body either by penetration through the wall of the food canal into coelomic cavities or by penetration into skeletal plates and spines. The spicules, which have penetrated into the coelom, obviously evoke a kind of protective answer. They were found to be entangled by clusters of cell remnants, the so-called brown bodies. The brown bodies contain melanin and gather at special sites of the echinoid body; these are the Stewart Organs, the gills and the inner side of the ambulacral plates. Sometimes the silicate spicule becomes surrounded by a calcareous sheath. The length of the sponge spicules makes their removal impossible, so that they are stored. The spicules penetrating into the plates are partly incorporated into the stereom. The four species examined in this study were Asthenosoma ijimai, Araeosoma owstoni, Diadema setosum (collected in Sagami Bay, Japan in 1991) and Hapalosoma gemmiferum (collected in Suruga Bay, Japan in 1991).     

Electron microscope study of the digestive system of Strongylocentrotus droebachiensis (Echinodermata: Echinoidea)

The fine structure of esophagus, stomach and intestinal epithelial cells of the sea urchin Strongylocentrotus droebachiensis have been examined. Three types of mucosal cells secreting neutral, acid and sulfated mucopolysaccharides have been found in the esophagus. The modes of origin of different types of mucous granules are described. Storage and zymogen cells have been identified in the stomach epithelium. Storage cells are adapted for intracellular digestion and synthesis of nutritive substance. Zymogen cells are adapted for enzyme synthesis. The columnar cells occur in the intestinal epithelium; they are adapted for nutrient absorption and lipid storage. Digestion in different parts of the digestive tract is considered in context with the ultrastructural and histochemical data obtained. Some problems concerning the relation between intracellular and extracellular digestion are discussed.     

Reproductive biology and biochemical composition of the brooding echinoid <Emphasis Type="Italic">Amphipneustes lorioli</Emphasis> on the Antarctic continental shelf

The bathyal West Antarctic Peninsula (WAP) shelf experiences intense seasonal variability in primary production, with summer phytoplankton blooms yielding intense pulse of phytodetritus to shelf sediments. Echinoderms form a conspicuous proportion of the deposit-feeding megabenthos on the shelf and of these Amphipneustes lorioli was the most abundant irregular echinoid. To explore the reproductive response of A. lorioli to this seasonal production cycle, A. Lorioli was sampled at one location on the WAP shelf during four separate cruises between March 2000 and March 2001. Reproductive patterns were determined by histological analyses of gonad tissue, and elemental (CHN) analyses were used to estimate the nutritional and energetic status of the body tissues. Histological analysis of the brooding echinoid A. lorioli suggested a quasi-continuous gametogenic pattern in both the ovaries and the testes. Biochemical analysis of the gonads and the gut tissues were consistent with a continuous gametogenic cycle, showing no significant changes in the biochemical composition of the tissues among seasons. Size-frequency distributions of the embryo and juvenile echinoids within the adults’ brood pouches revealed a synchronous recruitment of embryos and juveniles in specific cohorts between different adult specimens. Whilst this occurrence of different cohorts of the developing brood may be an adaptation to limited brood space, there may also be an external factor influencing the synchrony between adult individuals. Nonetheless, a continuous gametogenic cycle and the lack of seasonal variation in the biochemical composition of gonad and gut tissues suggest that this deposit-feeding irregular urchin is exploiting a persistent sediment food bank in WAP shelf sediments throughout much of the year.     

Particle capture in the musselMytilus edulis: The role of latero-frontal cirri

Microscope video graphs of particle paths near one-filament-thick mussel gill preparations, stimulated with a nerve transmitter (10^–6 M serotonin which restores normal ciliary activity), were used to disclose the capture of 6 m algal cells. Suspended algal cells carried with the water were stopped for a while at the entrance to the interfilament gap by the action of the latero-frontal cirri (Ifc), and transferred to the frontal side of the filament to be transported towards the marginal food groove. The event of transfer took place during approximately a time interval of 1150 to 1/25 s. To gain a better understanding of the capture mechanism and retention efficiency versus particle size, the flow through and around the Ifc was theoretically estimated. Normally beating Ifc create periodic, unsteady, three-dimensional flows at the entrance to the interfilament canal. During the active beat most of the water is deflected to flow around the branching cilia of the Ifc while some of the water is strained by these. Large particles (> 4 m) are stopped and transferred to the frontal current, whereas smaller particles either follow the flow around the Ifc and escape or they are stopped by the branching cilia.     

On endostyle ultrastructure in two new species of doliolid-like tunicates

The endostyle is a glandular and ciliated groove lining the pharyngeal floor of tunicates. This paper describes the histological structure of this organ in two new species of doliolid-like animals, Doliopsis bahamensis and Paradoliopsis harbisoni, and compares it with that of Doliolum nationalis. The results confirm the distinction between the Doliolida and the other Thaliacea. Received: 7 November 1996 / Accepted: 6 December 1996     

Animal-sediment relations in Cape Cod Bay,Massachusetts II. Reworking by Molpadia oolitica (Holothuroidea)

Populations of Molpadia oolitica (Holothuroidea) were sampled over an area of 440 km² in Cape Cod Bay, Massachusetts, USA. This species occurs in silt-clay muds in densities of 2 to 6/m², and is limited to depths greater than 22 m where the highest annual temperature does not exceed 10°C. M. oolitica lives within the mud, with its anterior end located approximately 20 cm below the surface. This species ingests only fine-grained particles at depth in the sediment, and deposits uncompacted feces at the surface, producing a fecal mound around the anal opening. This reworking produces vertical sediment sorting, high sediment-water content, and topographic relief of the sea floor. The fecal cones of M. oolitica provide a relatively stable surface for settlement and growth of the suspension feeders Euchone incolor (polychaete), Aeginina longicornis (amphipod), and Thyasira gouldi (bivalve) Uncompacted feces, accumulated in depressions between the mounds, form an unstable substratum frequently resuspended by tidal flow. Suspension feeders are absent from intercone areas.Contribution No. 236 of the Systematics-Ecology Program, Marine Biological Laboratory, Woods Hole, Massachusetts, USA.