Ultrastructure of the adhesive tentacles of the limnomedusa Vallentinia gabriella (Hydrozoa,Olindiadidae)

Summary The specialized adhesive exumbrellar tentacles of the limnomedusa Vallentinia gabriella were examined by light microscopy and scanning and transmission electron microscopy. The adhesive region first differentiates some distance from the tentacle tip. As differentiation proceeds the distal part is reduced and the adhesive region comes to lie at the tentacle tip. The adhesive epithelium consists of flagellated and non-flagellated glandular cells, a few nematocytes, and a nerve plexus. The glandular cells are characterized by electron-dense granules and bundles of microtubules. The microtubules, being anchored to the mesoglea, are oriented parallel to the longitudinal axis of the cell and extend up to the cell apex. It can be assumed that the microtubules are involved in the transport of secretory granules to the cell apex. Bundles of neurites run adjacent to the mesoglea between the basal processes of the glandular cells. The neurites form interneural synapses and synapses with glandular cells. It is suggested that detachment of the specialized adhesive tentacles is under nervous control.     

Secretory glands of the snail tentacle and their relation to the olfactory organ (Mollusca,Gastropoda)

Summary The epidermis of the posterior tentacles of the terrestrial snail Achatina fulica was examined by histological and histochemial methods. There are seven types of unicellular glands in the tentacle skin: three mucocytes containing either acid mucopolysaccharides or neutral mucopolysaccharides, or both; two mucocytes containing glycoproteins; a lipid gland; and a protein gland. The mucocytes are considerably more abundant along the shaft of the tentacle than at the tip, where the olfactory organ is situated. Conversely, the lipid glands and the protein glands are found almost exclusively in the olfactory organ. With minor exceptions, none of the foregoing cell types is present in the skin of the head or the foot. These observations indicate a high degree of local specificity in secretory products, consistent with a ubiquitous and generous endowment of glands in the molluscan skin. Collar cells, described by previous authors in closely related species, were not observed.     

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.     

Adhesive mechanisms in cephalopods: a review

Several genera of cephalopods (Nautilus, Sepia, Euprymna and Idiosepius) produce adhesive secretions, which are used for attachment to the substratum, for mating and to capture prey. These adhesive structures are located in different parts of the body, viz. in the digital tentacles (Nautilus), in the ventral surface of the mantle and fourth arm pair (Sepia), in the dorsal epidermis (Euprymna), or in the dorsal mantle side and partly on the fins (Idiosepius). Adhesion in Sepia is induced by suction of dermal structures on the mantle, while for Nautilus, Euprymna and Idiosepius adhesion is probably achieved by chemical substances. Histochemical studies indicate that in Nautilus and Idiosepius secretory cells that appear to be involved in adhesion stain for carbohydrates and protein, whilst in Euprymna only carbohydrates are detectable. De-adhesion is either achieved by muscle contraction of the tentacles and mantle (Nautilus and Sepia) or by secretion of substances (Euprymna). The de-adhesive mechanism used by Idiosepius remains unknown.     

Morphological,ultrastructural and histochemical investigation of epipodial sensory structures of Haliotis tuberculata (Gastropoda: Haliotidae)

In this study, we describe the microstructure and ultrastructure of the epipodial papillae and epipodial tentacles of Haliotis tuberculata using light and electron microscopy. The epipodial papillae vary morphologically; they are subdivided into several subpapillae whose surface is covered by small micropapillae. The epipodial tentacles are large extendable conically elongated structures whose surface is differentiated in two regions: the dorsal region with long corrugated folds, and a ventral region composed of three parts, a basal part with the same structure as the dorsal, a middle part with shorter corrugated folds and an apical part with large micropapillae. Although the thin sections and ultrastructure examination show that the epithelium of both organs is morphologically similar and composed of supporting cells, sensory cells and different types of secretory cells, there is a certain specialization in their secretory product. Although the epithelium of both structures was positive for acidic glycoconjugates, the tentacle epithelium was also positive for neutral sugars. Further specific differences were revealed by lectin histochemistry. Because papillae and tentacles can be extended or retracted depending on environmental conditions, they probably have tactile and olfactory functions.     

Fine Structure of the Tentacles of Phoronis australis Haswell (Phoronida,Lophophorata)

The epidermis of the tentacles of Phoronis australis consists of six cell types: supporting cells, choanocyte-like sensory cells, both types monociliated, secretory A-cells with a mucous secretion, and three kinds of B-cells with mucoprotein secretions. On cross-sections of the tentacle, one can distinguish four faces: the frontal one, heavily ciliated and located between the two frontolateral rows of sensory cells, the lateral and the abfrontal ones. The orientation of the basal structures of the cilia is related to the direction of their beat. The basiepidermal nervous system is grouped mainly at the frontal and abfrontal faces. The basement membrane is thickest on the frontal face and consists of circular collagen fibrils near the epidermis and longitudinal ones near the peritoneum. All peritoneal cells surrounding the mesocoel are provided with smooth longitudinal myofibrils, and isolated axons are situated between these cells and the basement membrane. The wall of the single blood capillary in each tentacle consists of epitheliomuscular cells with circular myofilaments, lying on a thin internal basal lamina; there is no endothelium.     

Ultrastructural study of two glandular systems in the proboscidial glandular epithelium of Riseriellus occultus (Nemertea, Heteronemertea)

 Two different types of glandular system in the proboscidial epithelium of Riseriellus occultus have been investigated by transmission electron microscopy. As expected, most of the epithelial cells are glandular in nature. With regard to differences in the ultrastructure of these gland cells and in the formation and morphology of their secretory granules, we have categorized and described four types of gland cell, indicated as G₁, G₂, G₃, and G₄. Each gland cell has a completely intraepithelial body characterized by a prominent nucleus, developed rough endoplasmic reticulum, Golgi complexes, and numerous secretory granules at different stages of maturation. These four types of gland cell appear associated in pairs forming numerous glandular systems of two types (A, B). These glandular systems are restricted to the ventral surface of the proboscis and are scattered irregularly throughout its length. Each glandular system consists of two gland cells of different types. The gland cell necks in each glandular system extend together to the epithelial surface; they protrude onto this and form a papilla where they open in a common area. The epithelial supportive cells adjacent to the glandular systems have long, stout microvilli which have a core of tonofilaments. These tonofilaments gather into dense bundles which pass vertically through the supportive cells and attach to the extracellular matrix underlaying the cells by hemidesmosomes. Moreover, a single sensory process stands close to each papilla. The ultrastructural morphology of the type A glandular systems suggests that they have an adhesive function operating in a similar way to that of the duo-gland adhesive systems in other invertebrate groups, although they are not homologous with these. The spatial arrangement of the secreted products of the type B glandular systems suggests that these may contribute to increasing the grip of the proboscis on the prey. The secretory granules (=pseudocnids) of the type G₃ gland cells are very likely an autapomorphy of the Anopla, providing a character by which the relationships within the Nemertea can be evaluated. Accepted: 9 October 1997     

The fine structure of the buccal tentacles of Holothuria forskali (Echinodermata,Holothuroidea)

Summary Ultrastructural study of the buccal tentacles of Holothuria forskali revealed that each tentacle bears numerous apical papillae. Each papilla consists of several differentiated sensory buds.The epidermis of the buds is composed of three cell types, i.e. mucus cells, ciliated cells, and glandular vesicular cells (GV cells). The GV cells have apical microvilli; they contain bundles of cross striated fibrillae associated with microtubules. Ciliated cells have a short non-motile cilium. Bud epidermal cells intimately contact an epineural nervous plate which is located slightly above the basement membrane of the epidermis. The epineural plate of each bud connects with the hyponeural nerve plexus of the tentacle. This nerve plexus consists of an axonic meshwork surrounded in places by sheath cells. The buccal tentacles have well-developed mesothelial muscles. Direct innervation of these muscles by the hyponeural nerve plexus was not seen.It is suggested that the buccal tentacles of H. forskali are sensory organs. They would recognize the organically richest areas of the sediment surface through the chemosensitive abilities of their apical buds. Tentacles presumably trap particles by wedging them between their buds and papillae.     

The gallbladder of the Capivara Hydrochoerus hydrochoeris (Mammalia: Rodentia)

The authors studied morphological and histochemically the mucopolysaccharides and proteins in the gallbladder tubular glands and epithelial cells of the capivara Hydrochoerus hydrochoeris. Based on the results the authors concluded: 1. the gallbladder single columnar epithelium consists of secretory, migrating, and goblet cells; 2. in the lamina propria are single coiled tubular glands; 3. goblet and tubular gland cells show neutral and sulphated mucopolysaccharides and sialic acid; 4. columnar cells show neutral mucopolysaccharides and protein radicals; 5. migrating cells show only protein radicals.     

The sensory epithelium of the tentacles and the rhinophore of Nautilus pompilius L. (cephalopoda, nautiloidea)

Nine intraepithelial ciliated cell types that are presumed to be sensory cells were identified in the epithelium of the pre- and postocular tentacles, the digital tentacles, and the rhinophore of the juvenile tetrabranchiate cephalopod Nautilus pompilius L. The morphological diversity and specialization in distribution of the different ciliated cell types analyzed by SEM methods suggest that these cells include receptors of several sensory functions. Ciliated cell types in different organs that show similar surface features were combined in named groups. The most striking cell, type I, is characterized by a tuft of long and numerous cilia. The highest density of this cell type occurs in ciliary fields in the epithelium of the lamellae of the pre- and postocular tentacles, in the olfactory pits of the rhinophores, and in the lamellae of four pairs of lateral digital tentacles, but not in the epithelium of the medial digital tentacles. The similar morphological data, together with behavioral observations on feeding habits, suggest that this cell type may serve in long-distance chemosensory function. The other ciliated cell types are solitary cells with specific spatial distributions in the various organs. Cell types with tufts of relatively short, stiff cilia (types III, IV, VIII), which are distributed in the lateral and aboral areas of the tentacles and at the base of the tentacle-like process of the rhinophore, are considered to be employed in mechanosensory transduction, while the solitary cells with bristle-like cilia at the margin of the ciliary fields (type II) and at the base of the rhinophore (type IX) may be involved in chemoreception. Histological investigation of the epithelium and the nerve structures of the different organs shows the proportion and distribution of the sensory pathways. Two different types of digital tentacles can be distinguished according to their putative functions: lateral slender digital tentacles in four pairs, of which the lowermost are the so-called long digital tentacles, participate in distance chemoreception, and the medial digital tentacles, whose terminal axial nerve cord may represent a specialized neuromechanosensory structure, appear to have contact chemoreceptive abilities.     

Fine Structure of Tentacles,Arms and Associated Coelomic Structures of Cephalodiscus gracilis (Pterobranchia,Hemichordata)

The tentacles of the pterobranch Cephalodiscus, a hemisessile ciliary feeder, originate from the lateral aspects of the arms and are covered by an innervated epithelium, the majority of its cells bearing microvilli. Each side of a tentacle has two rows of ciliated cells and additional glandular cells. The coelomic spaces in the tentacles are lined by cross-striated myoepithelial cells, allowing rapid movements of the tentacles. One, possibly two, blood vessels accompany the coelomic canal. On their outer sides the arms are covered by a simple ciliated epithelium with intra-epithelial nerve fibres; the inner side is covered by vacuolar cells. On both sides different types of exocrine cells occur. The collar canals of the mesocoel are of complicated structure. Ventrally their epithelium is pseudostratified and ciliated; dorsally it is lower and forms a fold with specialized cross-striated myoepithelial cells of the coelomic lining. Arms, tentacles, associated coelomic spaces and the collar canal of the mesocoel are considered to be functionally interrelated. It is assumed that rapid regulation of the pore width is possible and even necessary when the tentacular apparatus is retracted, which presumably leads to an increase of hydrostatic pressure in the coelom.     

Form and function of tentacles in pteriomorphian bivalves

Tentacles are remarkable anatomical structures in invertebrates for their diversity of form and function. In bivalves, tentacular organs are commonly associated with protective, secretory, and sensory roles. However, anatomical details are available for only a few species, rendering the diversity and evolution of bivalve tentacles still obscure. In Pteriomorphia, a clade including oysters, scallops, pearl oysters, and relatives, tentacles are abundant and diverse. We investigated tentacle anatomy in the group to understand variation, infer functions, and investigate patterns in tentacle diversity. Six species from four pteriomorphian families (Ostreidae, Pinnidae, Pteriidae, and Spondylidae) were collected and thoroughly investigated with integrative microscopy techniques, including histology, scanning electron microscopy, and confocal microscopy. Tentacles can be classified as middle fold tentacles (MFT) and inner fold tentacles (IFT) according to their position with respect to the folds of the mantle margin. While MFT morphology indicates intense secretion of mucosubstances, no evidence for secretory activity was found for IFT. However, both tentacle types have appropriate ciliary distribution and length to promote mucus transportation for cleaning and lubrication. Protective and sensory functions are discussed based on different lines of evidence, including secretion, cilia distribution, musculature, and innervation. Our results support the homology of MFT and IFT only for Pterioidea and Ostreoidea, considering their morphology, the presence of ciliated receptors at the tips, and branched innervation pattern. This is in accordance with recent phylogenetic hypotheses that support the close relationship between these superfamilies. In contrast, major structural differences indicate that MFT and IFT are probably not homologous across all pteriomorphians. By applying integrative microscopy, we were able to reveal anatomical elements that are essential for the understanding of homology and function when dealing with such superficially similar structures.     

Morphological and histological organization of the pyriform appendage of the tetrabranchiate Nautilus pompilius (Cephalopoda,Mollusca)

The pyriform appendage, an organ only found in nautiloid cephalopods was investigated with histological, histochemical and ultrastructural methods in order to characterize the anatomical and the cytological structure of this organ. The pyriform appendage is situated within the genital septum and lies in close contact with the ventricle of the heart. The proximal side ends blindly near the gonad whereas the distal side is developed into a duct. The duct was observed to open into the mantle cavity in juvenile and adult Nautilus pompilius of both sexes. Injections of India ink in the heart demonstrate that the organ is supplied with hemolymph from an artery that extends from the heart. The pyriform appendage is a hollow organ consisting mainly of glandular tissue. The lumen is covered with a columnar epithelium, the tunica mucosa, consisting of only one cell type containing vacuoles with different inclusions. Underneath the tunica mucosa is the tunica muscularis, which is embedded in connective tissue and folded, enlarging the internal surface. A cuboidal tunica serosa surrounds this organ. The vacuoles and the secretory products contain neutral mucopolysaccharides, glycoproteins and glycolipids. Acid phosphatase and serotonin were localized in the tunica mucosa. Acetylcholinesterase, catecholamines and the tetrapeptide FMRF‐amide were demonstrated within the nerve endings of the tunica muscularis indicating a dual “cholinergic‐aminergic” neuroregulation, possibly modulated by FMRF‐amide. These findings suggest that the pyriform appendage is not a rudimentary organ but instead has distinct biological functions in nautiloid cephalopods, possibly in intraspecific communication. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

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.     

Histology and histochemistry of the accessory reproductive glands in the male hairy-nosed wombat (Lasiorhinus latifrons)

The accessory male reproductive glands of the hairy-nosed wombat, Lasiorhinus latifrons, are a prostate and three pairs of Cowper's glands. Component units of all are branched tubular structures of varying epithelial makeup and secretory content. The prostate has the carrotlike shape and three consecutive regions commonly found in marsupials. The regions differ in their tubular histology and histochemistry: all contain secretory globules in glandular lumina. Cowper's glands A and B are histologically identical except for the absence of interstitial mast cells from gland G: gland C is characterized by narrower tubules and larger epithelial cells. Histochemical tests for protein, carbohydrate and iron indicate that glycogen is a major secretory product of the prostate (largely posterior region), iron is also secreted (mainly posterior region) and a small quantity of acid mucin is produced (mainly central region). Glycogen is a feature also of anterior prostatic glandular epithelium and of the capping cells of the urethral transitional epithelium. Cowper's gland A has considerable protein in its secretion, gland B a neutral glycoprotein and gland C a sialomucin: the latter two also exhibit cytoplasmic glycogen in their secretory cells.     

Comparative ultrastructure of catch tentacles and feeding tentacles in the sea anemone Haliplanella

TEM observations of catch tentacles revealed that the tentacle tip epidermis is filled with two size classes of mature holotrich nematocysts and a gland cell filled with electron-dense vesicles. Vesicle production is restricted to upper-middle and tentacle tip regions, whereas holotrich development occurs in the lower-middle and tentacle base regions. Thus, catch tentacles have a maturity gradient along their length, with mature tissues concentrated at the tentacle tip. Occasional feeding tentacle cnidae (microbasic p-mastigophores and basitrichs) and mucus gland cells occur in proximal portions of catch tentacles, but are phagocytized by amoeboid granulocytes and transported to the gastrodermis for further degradation. No feeding tentacle cnidae or mucus cells occur distally in catch tentacles. Unlike catch tentacles, feeding tentacles are homogeneous in structure along their length with enidocytes containing mature spirocysts, microbasic p-mastigophore or basitrich nematocysts distributed along the epithelial surface. Cnidoblasts are recessed beneath cnidocytes, occurring along the nerve plexus. Mucus gland cells and gland cells filled with electron-dense vesicles are present in feeding tentacles, distributed at the epithelial surface. Granular phagocytes are rare in the feeding tentacle tip, but common in the tentacle base.     

Histology and histochemistry of the accessory reproductive glands in the male hairy-nosed wombat (Lasiorhinus latifrons)

Summary The accessory male reproductive glands of the hairy-nosed wombat, Lasiorhinus latifrons, are a prostate and three pairs of Cowper's glands. Component units of all are branched tubular structures of varying epithelial makeup and secretory content. The prostate has the carrotlike shape and three consecutive regions commonly found in marsupials. The regions differ in their tubular histology and histochemistry: all contain secretory globules in glandular lumina. Cowper's glands A and B are histologically identical except for the absence of interstitial mast cells from gland B: gland C is characterized by narrower tubules and larger epithelial cells. Histochemical tests for protein, carbohydrate and iron indicate that glycogen is a major secretory product of the prostate (largely posterior region), iron is also secreted (mainly posterior region) and a small quantity of acid mucin is produced (mainly central region). Glycogen is a feature also of anterior prostatic glandular epithelium and of the capping cells of the urethral transitional epithelium. Cowper's gland A has considerable protein in its secretion, gland B a neutral glycoprotein and gland C a sialomucin: the latter two also exhibit cytoplasmic glycogen in their secretory cells.     

Evolutionary regularities of somatic polyploidy manifestation in salivary glands of gastropod molluscs. I. Subclasses Cyclobranchia and Scutibranchia

By means of histological and cytochemical methods, including DNA cytophotometry, the salivary glands of 11 species of molluscs of two old gastropod subclasses--Cyclobranchia and Scutibranchia (limpets)--have been investigated. In spite of some anatomical differences, the glandular epithelium of investigated molluscs includes functionally similar cell types: granular cells (with glycoproteid granular inclusions), mucocytes-I (that include sulfatic acid mucopolysaccharides), mucocytes-II (that include neutral and acid polysaccharides and proteins) and also the epithelial ciliated cells. Data of experiments on starvation and synchronous feeding of molluscs testify that all described cell types are independent. According to DNA cytophotometry data, the glandular cell nuclei are diploid in the main; only small part of the nuclei, varying in different species from 0.5 to 5.0%, displayed tetraploid DNA mass. A conclusion is made that in the oldest subclasses of gastropods (Cyclobranchia and Scutibranchia) somatic polyploidy, as a factor of tissue growth in salivary glands, is actually absent.     

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.