The ultrastructure of the tentacles of eleven species of dendrochirote holothurians studied with special reference to the surface coats and papillae

Summary The tentacles of eleven species of dendrochirote holothurians have been studied. The water vascular system, deep fibre system, ectoneural nerve ring and superficial fibre system are described and are similar to those of other holothurian tentacles. A conspicuous fuzzy coat covers the entire tentacular surface except for the tips of cilia. On the basis of its structure it is thought to be an attenuated glycocalyx. Its function is discussed in relation to anti-fouling and surface adhesion. The two surface coats underlying the fuzzy coat are termed the cuticle. Bacteria are found both within the surface coats and in the sub-cuticular space. Primary fixatives lacking osmium give poor preservation of the surface coats. The adhesive papillae of the apices of the tentacles contain elements of support cells and two other cells named Type-1 and Type-2 papillar cells. The secretions of Type-1 papillar cells are dense-cored vesicles and may contain a proteinaceous adhesive. The vesicles fuse with the cuticle and release their products which are apparently disseminated along the fuzzy coat filaments. The secretions of Type-2 papillar cells may have a neurosecretory function. The different models of food capture by dendrochirote tentacles are discussed as are duo-glandular adhesive systems in relation to dendrochirote tentacles.     

Ultrastructure of tentacles in the sipunculid worm <Emphasis Type="Italic">Thysanocardia nigra</Emphasis> Ikeda, 1904 (Sipuncula)

The ultrastructure of the tentacles was studied in the sipunculid worm Thysanocardia nigra. Flexible digitate tentacles are arranged into the dorsal and ventral tentacular crowns at the anterior end of the introvert of Th. nigra. The tentacle bears oral, lateral, and aboral rows of cilia; on the oral side, there is a longitudinal groove. Each tentacle contains two oral tentacular canals and an aboral tentacular canal. The oral side of the tentacle is covered by a simple columnar epithelium, which contains large glandular cells that secrete their products onto the apical surface of the epithelium. The lateral and aboral epithelia are composed of cuboidal and flattened cells. The tentacular canals are lined with a flattened coelomic epithelium that consists of podocytes with their processes and multiciliated cells. The tentacular canals are continuous with the radial coelomic canals of the head and constitute the terminal parts of the tentacular coelom, which shows a highly complex morphology. Five tentacular nerves and circular and longitudinal muscle bands lie in the connective tissue of the tentacle wall. Similarities and differences in the tentacle morphology between Th. nigra and other sipunculan species are discussed.Original Russian Text Copyright © 2005 by Biologiya Morya, Maiorova, Adrianov.     

Ultrastructure of the tentacles of Themiste lageniformis (Sipuncula)

Summary An ultrastructural study of the tentacles of Themiste lageniformis (Sipuncula) was conducted as part of a larger study of head metamorphosis in the species.The oral surface of the tentacles is constructed of a multiciliated, pseudostratified, columnar epithelium while the aboral surface is an unciliated, cuboidal epithelium. Intraepidermal mucous cells lie near the junction of the oral and aboral regions. The basal portion of the epidermal cells is embedded in a thick, collagenous extracellular matrix which contains outer circular muscles, inner longitudinal muscles, the main tentacular nerve and its branches. Three tentacular canals are present and are lined by peritoneum. Hemerythrocytes and coelomocytes flow through the lumen of the canals in a regular pattern.Justification for the designation of the tentacular canals as coelomic rather than vascular is discussed.     

Organization of the tentacular region in the vestimentiferan tubeworm <Emphasis Type="Italic">Riftia pachyptila</Emphasis> (Annelida,Vestimentifera)

Organization of the tentacular region in the giant vestimentiferan tubeworm Riftia pachyptila from hydrothermal vents has been reinvestigated. Protective cuticular structures consisting of a rod and a series of saucers have been found on the anterior surface of obturacula in juvenile individuals. This specific structural feature of juvenile R. pachyptila is regarded as a recapitulation of the ancestral state. The tentacular lamellae extend from the brain surface and run parallel to the body axis toward the anterior end of the tentacular region, which is similar to the structure of the tentacular apparatus in other vestimentiferan tubeworms. Its specific feature in R. pachyptila is the presence of vestimental extensions containing afferent and efferent blood vessels. Thus, differences in the structure of the tentacular region between R. pachyptila and other representatives of Tevniida are less than previously assumed, which allows R. pachyptila to be included in this group. It is considered that the specific structure of the tentacular region in R. pachyptila has developed as an adaptation allowing it to increase the number of tentacles.     

The fatty-acid composition and nutrition of deep-sea holothurians from the Sea of Okhotsk

The results of a comparative study of the fatty-acid composition in eight species of holothurians that were collected in the Sea of Okhotsk in the area of the Kuril Islands (depths of 90–560 m) are presented. It is shown that interspecific differences in the fatty-acid compositions of the holothurians were consistent with the isotope composition (δ¹³C and δ¹⁵N) and the structural features of the tentacles and the lifestyle of holothurians, as indicators of trophic resources used by these holothurians. According to the results of the cluster analysis, the holothurians were divided into three groups. The first group included suspension-feeding dendrochirotides Eupentacta pusilla and Pseudocnus fallax with a high content of the fatty acid 20:5(n-3), which is a marker of diatoms, and with the δ¹⁵N values that are typical of consumers of suspended organic matter. The second group consisted of the dendrochirotides Psolus chitonoides and Psolidium sp. with a much lower content of 20:5 (n-3) and higher contents of 20:4 (n-6) and 22:6 (n-3), as well as high values of δ¹⁵N, which are typical of surface deposit-feeders. The third group consisted of surface and subsurface depositfeeders, Chiridota sp., Molpadia orientalis, Pseudostichopus mollis, and Synallactes nozawai. Very high contents of 20: 4 (n-6) and 21: 4 (n-7) were typical of the third group and the highest values of δ¹⁵N, indicating feeding on repeatedly recycled organic matter.     

Ultrastructure of the tentacles of the apodous holothurian Leptosynapta spp (Holothurioidea: Echinodermata) with special reference to the epidermis and surface coats

Summary The tentacles of the apodous holothurian Genus Leptosynapta have been studied by use of transmission and scanning electron microscopy. The gross anatomy, water vascular system, fibre systems and ectoneural nerve ring are described. A fuzzy coat of attenuated filaments covers the surface of the tentacle, broken only by secretory ducts. A cuticle underlies the fuzzy coat. Bacteria are common in the subcuticular space. Fixation without osmium gives poor preservation of the surface coats. The epidermis consists of a single layer of columnar cells consisting of Type-1, Type-2, support, goblet and uniciliated cells. Type-1 cells secrete electron-dense material and appear to be homologous to adhesive cells of the tentacles of other holothurians. The support cells contain large, granular vesicles not found in other holothurians. Goblet cells contain flocculent mucus and have an apical cilium. Goblet cells are not found in other holothurian tentacles and may function to lubricate and wrap adhering particles to aid their ingestion. The uniciliated cells are rare, poorly developed and the cilium does not extend past the cuticle. The ultrastructure of the tentacles is discussed in relation to those of other holothurians.     

Ciliary feeding structures and particle capture mechanism in the freshwater bryozoan Plumatella repens (Phylactolaemata)

Abstract. In contrast to marine bryozoans, the lophophore structure and the ciliary filter‐feeding mechanism in freshwater bryozoans have so far been only poorly described. Specimens of the phylactolaemate bryozoan Plumatella repens were studied to clarify the tentacular ciliary structures and the particle capture mechanism. Scanning electron microscopy revealed that the tentacles of the lophophore have a frontal band of densely packed cilia, and on each side a zigzag row of laterofrontal cilia and a band of lateral cilia. Phalloidin‐linked fluorescent dye showed no sign of muscular tissue within the tentacles. Video microscopy was used to describe basic characteristics of particle capture. Suspended particles in the incoming water flow, set up by the lateral ‘pump’ cilia on the tentacles, approach the tentacles with a velocity of 1–2 mm s^‐1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve, as previously seen in marine bryozoans. The particle capture mechanism suggested is based on the assumed ability of the sensory stiff laterofrontal cilia to be triggered by the deflection caused by the drag force of the through‐flowing water on a captured food particle. Thus, when a particle is stopped by the laterofrontal cilia, the otherwise stiff cilia are presumably triggered to make an inward flick which brings the restrained particle back into the downward directed main current, possibly to be captured again further down in the lophophore before being carried to the mouth via the food groove. No tentacle flicks and no transport of captured particles on the frontal side of the tentacles were observed. The velocity of the metachronal wave of the water‐pumping lateral cilia was measured to be ～0.2 mm s^‐1, the wavelength was ～7 μm, and hence the ciliary beat frequency estimated to be ～30 Hz (～20 °C). The filter feeding process in P. repens reported here resembles the ciliary sieving process described for marine bryozoans in recent years, although no tentacle flicks were observed in P. repens. The phylogenetic position of the phylactolaemates is discussed in the light of these findings.     

Functional morphology of the tentacles and tentilla of Coeloplana bannworthi (Ctenophora, Platyctenida), an ectosymbiont of Diadema setosum (Echinodermata, Echinoida)

 The tentacular apparatus of Coeloplana bannworthi consists of a pair of tentacles which bear, on their ventral side, numerous tentilla. Each tentacle extends from and retracts into a tentacular sheath. Tentacles and tentilla are made up of an axial core covered by an epidermis. The epidermis includes six cell types: covering cells, two types of gland cells (mucous cells and granular gland cells), two types of sensory cells (ciliated cells and hoplocytes), and collocytes, this last cell type being exclusively found in the tentilla. The core is made up of a fibrillar matrix, the mesoglea, which is crossed by nerve processes and two kinds of smooth muscle cells. Regular muscle cells are present in both the tentacles and tentilla while giant muscle cells occur exclusively in the tentilla. The retraction of the tentacular apparatus is an active phenomenon due to the contraction of both types of muscle cells. The extension is a passive phenomenon that occurs when the muscle cells relax. Tentacles and tentilla first extend slightly due to the rebound elasticity of the mesogleal fibers and then drag forces exerted by the water column enable the tentacular apparatus to lengthen totally. Once the tentacles and tentilla are extended, gland cells, sensory cells, and collocytes are exposed to the water column. Any swimming planktonic organism may stimulate the sensory cilia which initiates tentillum movements. Pegs of hoplocytes can then more easily contact the prey which results in a slight elevation of the nearby collocytes, the last being responsible for gluing the prey to the tentilla. Accepted: 1 April 1997     

Tentacular diversity in deep-sea deposit-feeding holothurians: implications for biodiversity in the deep sea

The tentacles of deep-sea holothurians show a wide range of morphological diversity. The present paper examines gross tentacle morphology in surface deposit feeding holothurians from a range of bathymetric depths. Species studied included the elasipods: Oneirophanta mutabilis, Psychropotes longicauda and Benthogone rosea and the aspidochirotids: Paroriza prouhoi, Pseudostichopus sp., Bathyplotes natans and Paroriza pallens. The sympatric abyssal species Oneirophanta mutabilis, Psychropotes longicauda and Pseudostichopus sp. show subtle differences in diet and the structure and filling patterns of the gut that suggest differences in feeding strategies which may represent one mechanism to overcome competition for food resources in an environment where nutrient resources are considered to be, at least periodically, limiting. Interspecific differences in tentacle functional morphology and digestive strategies, which reflects taxonomic diversity could be explained in terms of Sanders'; Stability–Time Hypothesis. Since different tentacle types will turn over sediments to different extents, their impact on sedimentary communities will be enormous so that high diversity in meiofaunal communities may be explained most simply by Dayton and Hessler's Biological Disturbance Hypothesis.     

Endobacterial morphotypes in nudibranch cerata tips: a SEM analysis

The SEM investigation of nudibranch cerata material exhibits endobacterial morphotypes found in 12 out of 13 species tested: Aeolidia papillosa, Berghia caerulescens, Coryphella brownii, Coryphella lineata, Coryphella verrucosa, Cuthona amoena, Facelina coronata, Flabellina pedata, Dendronotus frondosus, Doto coronata, Tritonia plebeia and Janolus cristatus. Endobacteria could not be detected inside Tritonia hombergi. Endobacterial morphology found inside nudibranch species was compared to bacterial morphotypes detected earlier in tentacles of cnidarian species. SEM micrographs show endobacterial analogy among nudibranch species, but also similarity to cnidarian endobacteria investigated earlier. Of course, morphological data of microbes do not allow their identification. However, since most of these nudibranch species prey on cnidaria, it cannot be excluded that many of the endobacteria detected inside nudibranch species may originate from their cnidarian prey. Our previous data describing genetic affiliation of endobacteria from nudibranchian and cnidarian species support this assumption. Dominant coccoid endobacteria mostly exhibit smooth surface and are tightly packed as aggregates and/or wrapped in envelopes. Such bacterial aggregate type has been described previously in tentacles of the cnidarian species Sagartia elegans. Similar coccoid bacteria, lacking envelopes were also found in other nudibranch species. A different type of coccoid bacteria, characterized by a rough surface, was detected inside cerata of the nudibranch species Berghia caerulescens, and surprisingly, inside tentacles of the cnidarian species Tubularia indivisa. In contrast to cnidarian endobacteria, rod-shaped microorganisms are largely absent in nudibranch cerata.     

Tentacular apparatus ultrastructure in the larva of Bolinopsis infundibulum (Lobata: Ctenophora)

Borisenko, I. and Ereskovsky, A.V. 2011. Tentacular apparatus ultrastructure in the larva of Bolinopsis infundibulum (Lobata: Ctenophora). —Acta Zoologica (Stockholm) 00 : 1–10. Most ctenophores have a tentacular apparatus, which plays some role in their feeding. Tentacle structure has been described in adults of only three ctenophore species, but the larval tentacles have remained completely unstudied. We made a light and electron microscopic study of the tentacular apparatus in the larvae of Bolinopsis infundibulum from the White Sea. The tentacular apparatus of B. infundibulum larvae consists of the tentacle proper and the tentacle root. The former contains terminally differentiated cells, while the latter contains stem cells and cells undergoing differentiation. The core of the tentacle is formed by myocytes, and its epidermis contains colloblasts (hunting cells), wall cells, degenerating cask cells, refractive vesicles, and ciliated sensory cells. Stem cells, colloblasts, and cask cells at various stages of differentiation and putative myocytes progenitors were revealed in the tentacle root. Two different populations of the stem cells in the tentacle root give rise to epidermal (colloblasts and cask cells) and mesogleal (myocytes) cell lines. Nervous elements, glandular cells, and basal lamina were not found. Step‐by‐step differentiation of colloblasts and cask cells is described.     

Juvenile hormone induces ovarian development in diapausing cave-dwelling Drosophila species

Drosophila grisea and macroptera were collected in caves overwintering as adults. The females remained in a state of reproductive diapause which extended until May for macroptera and until July for grisea, whereas the males of both species had mature sperm at all times. Termination of the reproductive diapause under laboratory conditions was accomplished in grisea by exposing them to 14 hr of illumination daily and in macroptera by increasing the temperature to 20°C. Topical application of juvenile hormone (JH) on diapausing grisea caused a prompt termination of diapause and maturation of oöcytes within 10 days. Yolk proteins were found in the haemolymph of diapausing flies but not in their ovaries. In the JH-treated flies, yolk proteins were found in both the haemolymph and the ovaries, suggesting that in this species JH regulates the uptake of yolk proteins.     

Ultrastructure of the sensory epithelia of oral tube,fungiform sensory bodies,and terminal knobs of tentacles of Ovatella myosotis Draparnaud (Archaeopulmonata,Gastropoda)

Sensory epithelia of the oral tube, a fungiform body anterior to the tentacles and of the terminal knob of tentacles, were studied in Ovatella myosotis by electron microscopy. All three epithelia consist of columnar support cells, sensory cells, and, except in the oral tube, numerous goblet cells. The epithelia differ significantly in their apical differentiations. In the oral tube an outer layer is formed by irregularly bent villi of support cells completely embedded in a surface coat. Cilia and cytofila of the dendrites of sensory cells intertwine throughout the entire depth of the villous layer. In the fungiform sensory body some of the villi of support cells are singly branched. Their basal region is free of a surface coat. In this region cytofila and cilia of dendrites form a spongy layer, some cytofila extending into the surface coat. In the tentacular terminal knob the villi of the support cells branch dichotomously once or twice, a single villus thus ending with 2–4 tips. Only these terminal twigs are invested with the surface coat. The cytofila and dendritic cilia are confined to a broad spongy layer underneath. Three types of dendrites are present. They differ in their number of cilia, structure of basal bodies and occurrence in the three epithelia. Dendritic cytofila are most abundant in the tentacular terminal knob and least numerous in the oral tube. The observations are discussed with respect to corresponding epithelia in other pulmonates, the homology of the fungiform body, and possible functional correlates of structural features.     

Development of the tentacular apparatus in sipunculans (Sipuncula): I. Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920)

The development and arrangement of the tentacular apparatus of Thysanocardia nigra (Ikeda, 1904) and Themiste pyroides (Chamberlin, 1920) are described and illustrated using scanning electron microscopy. In T. nigra, the tentacular apparatus is composed of two crowns: the nuchal arc enclosing the nuchal organ and a crown of numerous oral tentacles arranged in U-shaped festoons. In early juveniles, two dorsal horn-like protrusions develop into the first, or primary, pair of tentacles of the nuchal arc. The second pair of tentacles of the nuchal arc develops dorsolaterally on the bases of the primary tentacles. Two ventrolateral lobes of the oral disk grow and become subdivided by the longitudinal ciliary groove into anlages of one set of dorsal and one set of ventral tentacles, thus forming a first oral festoon. Later, a pair of dorsolateral lobes develop between the first festoons and the nuchal arc to form a second pair of oral festoons. The third and following pairs of oral festoons develop in the dorsolateral growth zones lateral to the borders of the nuchal arc, where they meet the oral crown. The growing festoons extend down the oral disk and run alongside the head. A new oral tentacle appears directly at/on the base of the previous tentacle, thus giving rise to a typical sympodium with an alternating arrangement of tentacles. In T. pyroides, a second pair of tentacles develops from two ciliary lobes that are ventrolateral outgrowths of the circumoral ciliary field around the terminal mouth opening. The third pair of tentacles appears from the dorsolateral lobes at the base of primary tentacles, between the first two pairs of tentacles. These six tentacles determine the position of six main stems of the tentacular apparatus designated the first tentacles in the corresponding stems. The second tentacle in every stem appears as a ventrolateral outgrowth at the base of the first tentacle. The third and following tentacles in the stem are developed between the two previous tentacles according to a sympodial pattern. In both species, the distinct sympodial pattern in the arrangement of tentacles in the tentacular apparatus is well evidenced by the outlines of the ciliary oral grooves. The branched stems of T. pyroides may be homologized structurally and functionally to the oral festoons of T. nigra. J. Morphol. (c) 2006 Wiley-Liss, Inc.     

The morphology and life cycle of Ophryodendron mysidacii sp. nov. a marine suctorian epibiont on a mysid crustacean

A new species of suctorian protist epibiont of the mysid Schistomysis parkeri is described. The individuals show two types of adult form: elongated and flattened, both with 4-8 tentacular lobes. This new suctorian differs from described species of pro-Ophryodendron group by size, number of tentacular lobes, insertion of the tentacles, union of the lorica with the body, shape of the macronucleus, number of micronuclei and the lack of stalk (adult forms). The life cycle of this species is analysed and a succession pattern of its different stages is proposed.     

The thermotropic behavior and major molecular species composition of the phospholipids of echinoderms

This study examines the molecular species composition and heat-induced crystalline-liquid crystalline phase transitions of phosphatidylcholine (PC) and phosphatidylethanolamine (PE) from the muscle tissues of six species of echinoderms that were collected during the summer: the starfishes Distolasterias nipon and Asterias amurensis, the sea urchin Strongylocentrotus intermedius, and the holothurians Eupentacta fraudatrix, Cucumaria frondosa japonica, and Apostichopus japonicus. Phospholipids (PLs) were in the liquid crystalline state, which is optimal for the functioning of the cell membranes. The use of data on the molecular species composition of PLs for the interpretation of their thermotropic behavior indicated that homeoviscous adaptation is achieved by various rearrangements in the composition of the aliphatic groups of PLs. The phase transitions of PC and PE of echinoderms (except holothurians) were symbatic. The presence of a high-temperature peak on the PC thermograms of C. frondosa japonica and A. japonicus is attributable to the melting of the phospholipid domain, which is composed of molecular species with saturated aliphatic groups. Such domains are responsible for a significant shift in the temperature ranges of the phase transitions of phospholipids of holothurians and sea urchin towards temperatures above 0°C.     

The feeding behaviour of a deep-sea holothurian, Stichopus tremulus (Gunnerus) based on in situ observations and experiments using a Remotely Operated Vehicle

Using a Remotely Operated Vehicle (ROV) to deploy an in situ cage experiment incorporating fluorescent Luminophore particle tracers, the gut throughput time of the deposit feeding holothurian, Stichopus tremulus (Gunnerus) was determined as 23.73 h (S.D.±2.3). For a range of individuals examined at different depths (350-500 m) and locations, throughput times varied between 19 and 26 h irrespective of animal size or gut tract length. In situ video observations of feeding behaviour showed that this species uses fine oral papillae in a ‘sweeping’ motion to target particles on the seafloor. Following detection of a food source fine-branched digitate tentacles collect a large range of sediment fragments from the seabed. The main types of particles ingested include silica fragments (<20 >500 μm), pelagic foraminifera, benthic foraminifera, fine phytodetrital remains and occasional larger rock fragments (∼1 cm). Ingested sediment consisted mainly of very fine silica fragments (∼50 μm) accounting for over 50% of the total gut contents. Frame-by-frame video analysis revealed that the particle handling time (i.e. the time taken for a tentacle insertion and the subsequent collection of food) was found to be ∼54 s. Only 10 of the 20 feeding tentacles were simultaneously employed during feeding. Use of tentacles appeared to be in sequence, alternating between the reserve and active tentacles. Estimating the rate of movement over the seabed and the total effective capture area of each tentacle, the impact of this animal on the turnover and quality of surface sediment at this deepwater site is potentially substantial. The in situ experiments provided a significant improvement over previous methods used to investigate deep-sea deposit feeders and represent a useful concept for further in situ deep-sea research using an industrial ROV.     

Elektronenmikroskopische Untersuchungen an den Oralcirren und der Haut vonBranchiostoma lanceolatum

Praeoral tentacles and epidermis of the anterior body region ofBranchiostoma lanceolatum Pallas have been investigated by electron microscopy. The epidermis of the praeoral tentacles and the anterior body region are mono-layered and cohere by strong denticulations of the adjoining cell walls. Vertical secretory vesicles at the cell surface give off mucous substances. The secretory vesicles are found only in the body epithelium. Between epithelium cells both epithelia contain two different secondary sensilla types.B. lanceolatum is the lowest chordate in which taste buds of the praeoral tentacles have been found. The taste buds overtop the surface of the epithelium. The praeoral tentacles are stiffened by a skeleton rod, situated asymmetrically and build up in layers. The skeleton rod is surrounded by connective tissue, which includes a coelomic space. Axon bundles of different strength are situated in the connective tissue. Not only the taste buds but also singular sensilla types are innervated by these axon bundles. The relatively strong basement lamina is partially zonated and contains pores. An antagonistically arranged layer of collagen fibres of varying thickness occurs below the basement lamina.     

Structural and Developmental Disparity in the Tentacles of the Moon Jellyfish Aurelia sp.1

Tentacles armed with stinging cells (cnidocytes) are a defining trait of the cnidarians, a phylum that includes sea anemones, corals, jellyfish, and hydras. While cnidarian tentacles are generally characterized as structures evolved for feeding and defense, significant variation exists between the tentacles of different species, and within the same species across different life stages and/or body regions. Such diversity suggests cryptic distinctions exist in tentacle function. In this paper, we use confocal and transmission electron microscopy to contrast the structure and development of tentacles in the moon jellyfish, Aurelia species 1. We show that polyp oral tentacles and medusa marginal tentacles display markedly different cellular and muscular architecture, as well as distinct patterns of cellular proliferation during growth. Many structural differences between these tentacle types may reflect biomechanical solutions to different feeding strategies, although further work would be required for a precise mechanistic understanding. However, differences in cell proliferation dynamics suggests that the two tentacle forms lack a conserved mechanism of development, challenging the textbook-notion that cnidarian tentacles can be homologized into a conserved bauplan.