Ultrastructural study of the jaw structures in two species of Ampharetidae (Annelida: Polychaeta)

Two species of jaw bearing Ampharetidae (Adercodon pleijeli (Mackie 1994) and Ampharete sp. B) were investigated in order to describe the microanatomy of the mouth parts and especially jaws of these enigmatic polychaetes. The animals of both studied species have 14–18 mouth tentacles that are about 30 µm in diameter each. In both species, the ventral pharyngeal organ is well developed and situated on the ventral side of the buccal cavity. It is composed of a ventral muscle bulb and investing muscles. The bulb consists of posterior and anterior parts separated by a deep median transversal groove. In both species, the triangular teeth or denticles are arranged in a single transversal row on the surface of the posterior part of the ventral bulb just in front of its posterior edge. There are 36 denticles in Adercodon pleijeli and 50 in Ampharete sp. B. The height of the denticles (6–12 µm) is similar in both species. Each tooth is composed of two main layers. The outer one (dental) is the electron‐dense sclerotized layer that covers the tooth. The inner one consists of long microvilli with a collagen matrix between them. The thickness of the dental layer ranges from 0.95 to 0.6 µm. The jaws of the studied worms may play a certain role in scraping off microfouling. The fine structure of the jaws in Ampharetidae is very similar to that of the mandibles of Dorvilleidae, the mandibles and the maxillae of Lumbrineridae, Eunicidae and Onuphidae, and the jaws of other Aciculata. This type of jaw is characterized by unlimited growth and the absence of replacement. The occurrence of jaws in a few smaller Ampharetidae is considered as an apomorphic state.     

Morphology of the jaw apparatus in 8 species of Patellogastropoda (Mollusca, Gastropoda) with special reference to Testudinalia tesulata (Lottiidae)

The fine structure of the jaw apparatus was studied by scanning electron microscopy in eight species of Patellogastropoda. The jaw apparatus is an unpaired two-layered dorsolateral structure with anterior and posterior wings attached to the odontophore by muscles. The jaw of Testudinalia tesulata (O.F. Müller, 1776) is a derivative of the cuticle typical for the foregut. The tissue forming the jaw is a specialized foregut epithelium (gnathoepithelium), consisting of a special type of cells called gnathoblasts. The jaw grows in areas of the epithelium characterized by high concentration of electron-dense vesicles, ER and long microvilli that penetrate deep into the jaw plate. This indicates that the gnathoblasts take an active part in jaw growth. In most cases, these areas of the gnathoepithelium are highly folded. The main differences between the species studied are form and thickness of the frontal edge of the jaw. These differences do not correlate with the systematic position of the species studied but likely depend more on the feeding mode. The transmission electron microscopy studies yielded new morphological criteria for comparison between various gastropod species and other members of Trochozoa, in particular, Annelida. The jaws of Annelida are cuticular structures formed on the surface of specialized epithelial cells, often also called gnathoblasts. The jaw of Patellogastropoda can be attributed to the first type of annelid jaw formation characterized by an epithelium with long microvilli and continuous growth.     

Eating with a saw for a jaw: Functional morphology of the jaws and tooth‐whorl in Helicoprion davisii

The recent reexamination of a tooth‐whorl fossil of Helicoprion containing intact jaws shows that the symphyseal tooth‐whorl occupies the entire length of Meckel's cartilage. Here, we use the morphology of the jaws and tooth‐whorl to reconstruct the jaw musculature and develop a biomechanical model of the feeding mechanism in these early Permian predators. The jaw muscles may have generated large bite‐forces; however, the mechanics of the jaws and whorl suggest that Helicoprion was better equipped for feeding on soft‐bodied prey. Hard shelled prey would tend to slip anteriorly from the closing jaws due to the curvature of the tooth‐whorl, lack of cuspate teeth on the palatoquadrate (PQ), and resistance of the prey. When feeding on soft‐bodied prey, deformation of the prey traps prey tissue between the two halves of the PQ and the whorl. The curvature of the tooth‐whorl and position of the exposed teeth relative to the jaw joint results in multiple tooth functions from anterior to posterior tooth that aid in feeding on soft‐bodied prey. Posterior teeth cut and push prey deeper into the oral cavity, while middle teeth pierce and cut, and anterior teeth hook and drag more of the prey into the mouth. Furthermore, the anterior‐posterior edges of the teeth facilitate prey cutting with jaw closure and jaw depression. The paths traveled by each tooth during jaw depression are reminiscent of curved pathways used with slashing weaponry such as swords and knifes. Thus, the jaws and tooth‐whorl may have formed a multifunctional tool for capturing, processing, and transporting prey by cyclic opening and closing of the lower jaw in a sawing fashion. J. Morphol. 276:47–64, 2015. © 2014 Wiley Periodicals, Inc.     

Ultrastructure and function of the pharynx of Gnathostomula paradoxa (Gnathostomulida)

 The pharynx of Gnathostomula paradoxa consists of the partly syncytial pharyngeal musculature, a pharyngeal epithelium, myoepitheliocytes, receptors, nerves, and three solid parts, called the jugum, the basal plate, and the jaw. Extended non-contractile regions of both pharyngeal and body wall musculature form the so-called parenchymatous tissue between the digestive tract and the body wall. The pharyngeal epithelium mediates the force from the pharyngeal musculature to the solid parts. The basal plate and jaw contain longitudinal cuticular rods which are elastic antagonists of the musculature. There is no buccal ganglion in G. paradoxa. The study supports the monophyly of the Gnathostomulida and Gnathifera. Accepted: 4 April 1997     

Six Hydrophobins Are Involved in Hydrophobin Rodlet Formation in Aspergillus nidulans and Contribute to Hydrophobicity of the Spore Surface

Hydrophobins are amphiphilic proteins able to self-assemble at water-air interphases and are only found in filamentous fungi. In Aspergillus nidulans two hydrophobins, RodA and DewA, have been characterized, which both localize on the conidiospore surface and contribute to its hydrophobicity. RodA is the constituent protein of very regularly arranged rodlets, 10 nm in diameter. Here we analyzed four more hydrophobins, DewB-E, in A. nidulans and found that all six hydrophobins contribute to the hydrophobic surface of the conidiospores but only deletion of rodA caused loss of the rodlet structure. Analysis of the rodlets in the dewB-E deletion strains with atomic force microscopy revealed that the rodlets appeared less robust. Expression of DewA and DewB driven from the rodA promoter and secreted with the RodA secretion signal in a strain lacking RodA, restored partly the hydrophobicity. DewA and B were able to form rodlets to some extent but never reached the rodlet structure of RodA. The rodlet-lacking rodA-deletion strain opens the possibility to systematically study rodlet formation of other natural or synthetic hydrophobins.     

Characteristics of Streptomyces coelicolor A3(2) aerial spore rodlet mosaic

Cytochemical analysis of Streptomyces coelicolor (A3(2) indicated that the aerial growth rodlet mosaic is a polysaccharide. Statistical analysis of frequency distributions of individual rodlet lengths from control and ether-reoriented spore mosaics indicated that the rodlet fibrillar image is the result of individual particulates, rather than evaginations in a continuous sheet of material. A model of the mature sport envelope was developed from freeze-etch-replicated, thin-sectioned, and critical point dried S. coelicolor A3(2) mature spores. The rodlet mosaic was situated between the outer spore wall and an external granuloma matrix. Mixture spore envelope layers from the inner surface to the external surface are plasma membrane, inner spore wall, outer spore wall, rodlet mosaic, an undefined granular matrix, and the sheath. The granular matrix had an uneven thickness and much of the matrix was frequently absent from the interspore spaces of mature spore chains. Streptomyces coelicolor A3(2) mosaic rodlets were isolated by acetic acid refluxing, then ethanol precipitation. Complete acid hydrolysis of rodlets released on sugar which cochromatographed with D-glucosamine-HCl and released acetic acid at 139% of the expected level. Cell associated rodlet mosaics and isolated mosaic rodlets were hydrolyzed with chitinase. Infrared spectra of isolated rodlets were similar to crab chitin spectra.     

Self-assembly of proteins into a three-dimensional multilayer system: Investigation of the surface of the human fungal pathogen Aspergillus fumigatus

Hydrophobins are small surface active proteins that fulfil a wide spectrum of functions in fungal growth and development. The human fungal pathogen Aspergillus fumigatus expresses RodA hydrophobins that self-assemble on the outer conidial surface into tightly organized nanorods known as rodlets. AFM investigation of the conidial surface allows us to evidence that RodA hydrophobins self-assemble into rodlets through bilayers. Within bilayers, hydrophilic domains of hydrophobins point inward, thus making a hydrophilic core, while hydrophobic domains point outward. AFM measurements reveal that several rodlet bilayers are present on the conidial surface thus showing that proteins self-assemble into a complex three-dimensional multilayer system. The self-assembly of RodA hydrophobins into rodlets results from attractive interactions between stacked β-sheets, which conduct to a final linear cross-β spine structure. A Monte Carlo simulation shows that anisotropic interactions are the main driving forces leading the hydrophobins to self-assemble into parallel rodlets, which are further structured in nanodomains. Taken together, these findings allow us to propose a mechanism, which conducts RodA hydrophobins to a highly ordered rodlet structure. The mechanism of hydrophobin assembly into rodlets offers new prospects for the development of more efficient strategies leading to disruption of rodlet formation allowing a rapid detection of the fungus by the immune system.     

Chemical composition and electron microscopy of the rodlet layer ofAspergillus nidulans conidia

The rodlet layers of wild-type and white mutantAspergillus nidulans conidia were purified by a simple centrifugation procedure after conidial suspensions were subjected to sonication. Chemical analysis showed that the major components of wild-type rodlets were protein and melanin in almost equal amounts, followed by carbohydrate. White mutant rodlets differed from those of the wild-type strain in that the melanin content was very low. Histidine, aspartic acid, glutamic acid, glycine, and alanine were the most prominent amino acids in the rodlet layer of the white mutant, whereas cystine and methionine were not found. Electron microscopy studies showed that the rodlets of both white mutant and wild-type strains were grouped into fascicles, which varied from 80 to 160 nm in width. Individual rodlets measured 125–360 nm in length and 7 nm in diameter.     

The Cys3-Cys4 loop of the hydrophobin EAS is not required for rodlet formation and surface activity

Class I hydrophobins are fungal proteins that self-assemble into robust amphipathic rodlet monolayers on the surface of aerial structures such as spores and fruiting bodies. These layers share many structural characteristics with amyloid fibrils and belong to the growing family of functional amyloid-like materials produced by microorganisms. Although the three-dimensional structure of the soluble monomeric form of a class I hydrophobin has been determined, little is known about the molecular structure of the rodlets or their assembly mechanism. Several models have been proposed, some of which suggest that the Cys3-Cys4 loop has a critical role in the initiation of assembly or in the polymeric structure. In order to provide insight into the relationship between hydrophobin sequence and rodlet assembly, we investigated the role of the Cys3-Cys4 loop in EAS, a class I hydrophobin from Neurospora crassa. Remarkably, deletion of up to 15 residues from this 25-residue loop does not impair rodlet formation or reduce the surface activity of the protein, and the physicochemical properties of rodlets formed by this mutant are indistinguishable from those of its full-length counterpart. In addition, the core structure of the truncation mutant is essentially unchanged. Molecular dynamics simulations carried out on the full-length protein and this truncation mutant binding to an air-water interface show that, although it is hydrophobic, the loop does not play a role in positioning the protein at the surface. These results demonstrate that the Cys3-Cys4 loop does not have an integral role in the formation or structure of the rodlets and that the major determinant of the unique properties of these proteins is the amphipathic core structure, which is likely to be preserved in all hydrophobins despite the high degree of sequence variation across the family.     

Tooth growth and replacement in Ctenolucius hujeta, a neotropical characoid fish

The predaceous neotropical characoid fish Ctenolucius has an essentially homodont dentition, the number of teeth increasing linearly with age. The basic manner of tooth replacement suggests that Ctenolucius is a primitive characoid. Tooth replacement continues throughout life and is similar to that of tetrapods, involving replacement waves which pass from the back to the front of the jaws. The waves containing the greatest number of teeth are found just anterior to the middle of the jaws. In the upper jaw the increase in the number of teeth is restricted to the anterior portion (premaxillary) whereas the number on the posterior part (maxillary) remains constant. In specimens measuring from 68–230 mm in standard length the posterior portion of the upper jaw doubles in length whereas the anterior portion triples. It is suggested that the area immediately anterior to the middle of the jaw, where replacement waves are longest, is where most of the increase in tooth numbers occurs. During growth of the teeth the absolute height is always greater than the absolute width as the shape changes. The final shape of the recurved conical teeth is determined only in the last stages of tooth formation when the main axis of growth abruptly changes.     

Anatomy and Ultrastructure of Ventral Pharyngeal Organs and their Phylogenetic Importance in Polychaeta (Annelida). IV. The Pharynx and Jaws of the Dorvilleidae

An anatomical and ultrastructural investigation of the ventral pharyngeal organ, jaws and replacement of jaws was carried out in Ophryotrocha gracilis and Protodorvillea kefersteini (Dorvilleidae). The pharynx exhibits the following features: jaw apparatus present, consisting of paired mandibles and rows of maxillary plates, the latter are fused to form a single piece; cuticular jaws electron-dense, in P. kefersteini with collagen fibres; muscle bulbus solid, composed of muscle cells only; parallel running myofilaments, centrally located mitochondria and nuclei, bulbus epithelium containing the mandibles and gland cells, maxillary plates lying on folds corresponding to a tongue-like organ, connected with mandibles by longitudinal investing muscles; numerous gland cells not united to distinct salivary glands. Development of jaw replacements occurs in epithelial cavities beside the functional maxillae. Shape of maxillary plates is preformed by microvilli carrying cell processes. Maxilloblasts change their shape during the development. Synapomorphic structures occurring in ventral pharyngeal organs of other species outside the Eunicea are not present and even the closely related Dinophilidae exhibit a completely different pharyngeal organ. Therefore, convergent evolution of these organs is the most probable explanation. These findings do not agree with the hypothesis of the homology of the ventral pharyngeal organs in the Polychaeta.     

First record ofColoborhynchus (Pterosauria) from the Santana Formation (Lower Cretaceous) of the Chapada do Araripe, Brazil

The anterior tips of associated upper and lower jaws of a pterosaur from the Lower Cretaceous of Brazil are described and assigned to the taxonColoborhynchus in the family Ornithocheiridae. It is characterized by the shape and position of the sagittal crest on the upper and lower jaw, the arrangement and length of the teeth and the spoon-like lateral expansion of the anterior parts of the jaws. It closely resemblesColoborhynchus wadleighi from North America andColoborhynchus clavirostris from England. Diagnostic anatomical characteristics permit a revision of the genusTropeognathus, which is shown here to be a junior synonym of other described taxa.Tropeognathus mesembrinus is referred toCriorhynchus andT. robustus toColoboryhnchus. Consistent anatomical features enable the new jaw fragment to be assigned toColoborhynchus robustus.      

Morphological and functional bases of durophagy in the queen triggerfish,Balistes vetula (Pisces,tetraodontiformes)

Tetraodontiform fishes are characterized by jaws specialized for powerful biting and a diet dominated by hard-shelled prey. Strong biting by the oral jaws is an unusual feature among teleosts. We present a functional morphological analysis of the feeding mechanism of a representative tetraodontiform, Balistes vetula. As is typical for the order, long, sharp, strong teeth are mounted on the short, robust jaw bones of B. vetula. The neurocranium and suspensorium are enlarged and strengthened to serve as sites of attachment for the greatly hypertrophied adductor mandibulae muscles. Electromyographic recordings made from 11 cranial muscles during feeding revealed four distinct behaviors in the feeding repertoire of B. vetula. Suction is used effectively to capture soft prey and is associated with a motor pattern similar to that reported for many other teleosts. However, when feeding on hard prey, B. vetula directly bit the prey, exhibiting a motor pattern very different from that of suction feeding. During buccal manipulation, repeated cycles of jaw opening and closing (biting) were coupled with rapid movement of the prey in and out of the mouth. Muscle activity during buccal manipulation was similar to that seen during bite-captures. A blowing behavior was periodically employed during prey handling, as prey were forcefully “spit out” from the mouth, either to reposition them or to separate unwanted material from flesh. The motor pattern used during blowing was distinct from similar behaviors described for other fishes, indicating that this behaviors may be unique to tetraodontiforms. Thus B. vetula combines primitive behaviors and motor patterns (suction feeding and buccal manipulation) with specialized morphology (strong teeth, robust jaws, and hypertrophied adductor muscles) and a novel behavior (blowing) to exploit armored prey such as sea urchins molluscs, and crabs. © 1993 Wiley-Liss, Inc.     

Observations on rodlet cells found in the vascular system and extravascular space of angelfish (Pterophyllum scalare scalare)

In the angelfish ( Pterophyllum scalare scalare ) numerous rodlet cells were found in the large post-orbital blood vessel caudal to the eye and in the surrounding extravascular space. Within the vessel the rodlet cells formed striking regular arrays, along the inner aspect of the wall. The rodlets within the cells were positive to PAS but negative to Sudan Black B, Masson's, and the Fuelgen stain. The capsule around the cells was negative for all these stains. These rodlet cells appeared to be traversing the vessel endothelium, and to be pushing the endothelium aside without damaging it. Some discharged their contents into the vessel, but we never observed the release of intact rodlets. The nuclei of rodlet cells in actual contact with the vessel were at the end of the cell more distant from the endothelial wall. Cell-to-cell adhesion structures or communications junctions between rodlet cells and the endothelium were not evident. A putative rodlet cell precursor in the extravascular space contained large electron-dense granules, and extended pseudopodia that contacted nearby rodlet cells. Based on their morphology, tissue distribution, and their behaviour, we conclude that the rodlet cell is an endogeneous teleost cell type, and possibly represents a form of matured granulocyte.     

Development of rodlet cells in the gut of turbot (Psetta maxima L.): Relationship between their morphology and S100 protein immunoreactivity

Rodlet cells are an enigmatic cell type described in tissues of both marine and freshwater teleosts. Although their structure is well established, up to date their function remains subject of debate. However, there is consensus among the majority of researchers that rodlet cells play an important role within immune system, and this function is probably related with the release of rodlets due to contractile capability of their fibrous layer. Regulation of the contraction mechanism would require proteins that modulate Ca⁺⁺ intracellular concentration to be expressed in rodlet cells. We performed a morphological and immunohistochemical study at light and electron microscopy levels to assess S100 protein immunoreactivity in developing rodlet cells. Immature stages did not exhibit immunoreactive signal; however, immunoreactivity was observed in the fibrous layer of both transitional and mature rodlet cells. The latter stage also showed immunosignal within the rodlets. These findings suggest a clear association between S100 protein expression and rodlet cell development that could be linked to the regulation of rodlet activity and contractile property of their fibrous layer. Furthermore, S100 protein antibody constitutes a novel marker for rodlet cells that could be used in future studies of this particular cell type.     

Recruitment of class I hydrophobins to the air:water interface initiates a multi-step process of functional amyloid formation

Class I fungal hydrophobins form amphipathic monolayers composed of amyloid rodlets. This is a remarkable case of functional amyloid formation in that a hydrophobic:hydrophilic interface is required to trigger the self-assembly of the proteins. The mechanism of rodlet formation and the role of the interface in this process have not been well understood. Here, we have studied the effect of a range of additives, including ionic liquids, alcohols, and detergents, on rodlet formation by two class I hydrophobins, EAS and DewA. Although the conformation of the hydrophobins in these different solutions is not altered, we observe that the rate of rodlet formation is slowed as the surface tension of the solution is decreased, regardless of the nature of the additive. These results suggest that interface properties are of critical importance for the recruitment, alignment, and structural rearrangement of the amphipathic hydrophobin monomers. This work gives insight into the forces that drive macromolecular assembly of this unique family of proteins and allows us to propose a three-stage model for the interface-driven formation of rodlets.     

Tenuignathia rikerae nov. gen., nov. spec., a New Gnathostomulid from the West Atlantic

A further paper dealing with West Atlantic Gnathostomulida presents the new monotypic genus Tenuignathia. The presence of a full sensorium and a cuticular bursa place the new genus within the order Bursovaginoidea, sub-order Scleroperalia; the simple jaws and pharynx, the bipartite ciliary pits as well as epithelial characters clearly suggest inclusion in the family Mesognathariidae. Found in Florida, North Carolina and Bermuda so far, Tenuignathia displays an unusually wide range of jaw length even within a narrow geographic range, which will be the subject of a separate study.     

An ultrastructural approach to feeding behaviour in Philodina roseola and Brachionus calyciflorus (rotifers) II. The oesophagus

The cuticular oesophagus is a simple expansion of the dorsal pharyngeal wall of the mastax. The ciliary oesophagus is the cellular anterior wall of the stomach lumen, but seems to have the same embryological origin as the pharynx.In Brachionus calyciflorus, its cilia are surrounded by cuticular velums which have the same myelin-like structure and the same function as the buccal velum of Philodina roseola. In all cases, the oesophagus prevents the return of food particles from the stomach to the mastax lumen.     

Experimental and comparative morphology of radula renewal in pulmonates (Mollusca,Gastropoda)

Summary The continuous renewal of the pulmonate radula and the histology and regeneration of its concomitant epithelia were studied by light and electron microscopy, autoradiography and electron microprobe analysis. The two species investigated show histological differences and the results were compared with those of a preceding study on a prosobranch radula. The radula is an intricate cuticular structure of the foregut. Only the fully grown part, which is active during feeding, lies in the buccal cavity while it is constantly renewed by the coordinated cooperation of specialized cells forming the radular sheath. The end of the sheath is occupied by cells which produce the organic matrix of the radula. In taeniogloss prosobranchs, seven multicellular cushions of small odontoblasts lie at the end of the sheath and produce the seven teeth of each cross-row. In pulmonates, the multidenticular radula is generated by numerous groups of a few voluminuous cells. Despite these histological differences, prosobranchs and pulmonates generate the radula matrix by microvilli, cytoplasmatic protrusions and apocrine secretions. The epithelia of the radular sheath contribute to the transport, tanning and mineralization of the radula. The concomitant epithelia are replaced in limited proliferation zones at the end of the radular sheath and their cells migrate anteriorly to the buccal cavity. The ultrastructure of the sheath cells and the alterations which they undergo in connection with their functions are discussed. The proliferation zone of the superior epithelium is strictly confined and the cells move together with the radula forward. In prosobranchs, the cells of the superior epithelium begin to degenerate in the middle of the radular sheath and the entire epithelium is simply extruded into the buccal cavity. In pulmonates, the opening of the radular sheath is closed by the cuticular collostylar hood which is generated by a distinct epithelium which is proved to be stationary. When leaving the proliferation zone, the superior epithelium differentiates into supporting cells and mineralizing cells; the latter cause the hardening of the radular teeth and already degenerate in the middle of the sheath. The whole superior epithelium degenerates at the border to the collostylar hood-epithelium. In Lymnaea the degeneration zone is strictly confined whereas in Cepaea the collostylar hood and its generating epithelium extend into the radular sheath and the degeneration zone ranges over a distance of 3–5 rows of teeth. The proliferation zone of the inferior epithelium extends over the posterior half of the radular sheath, but the replacement rate is much lower than in the superior epithelium. Although the inferior epithelium carries the radula, it migrates slower than the radula. Obviously the radula has to be transported actively by apical protrusions of the cells, which penetrate into the radular membrane. At the opening of the radular sheath the inferior epithelium generates the adhesive layer and degenerates. During feeding, the adhesive layer has to maintain the firm mechanical connection between radula and distal radular epithelium. Autoradiographic experiments demonstrate that the distal radular epithelium is stationary. Nevertheless, the radula is known to advance to its degeneration zone. Special attention is paid to this problem. We strongly suspect that the transport of the adhesive layer and the radula is based on pseudopodial movements of apical protrusions characteristic for the distal radular epithelium. These protrusions interdigitate with the lower face of the adhesive layer. The mechanical connection has to be maintained and so the respective structures (tonofilaments and hemi-desmosomes) have to be continually renewed. This needs a high amount of energy and obviously results in the conspicuous concentration of mitochondria near the apical surface.Abbreviations al adhesive layer - ax axon - bc buccal cavity - bce buccal cavity epithelium - bl basal layer - bla basal labyrinth - bm basal membrane - bp basal plate - bpc basal plate cell - c cilia - ch collostylar hood - che collostylar hood-epithelium - cl cuticular layer - col collostyle - cr cell remnant - cts connective tissue sheath - d desmosome - dl upper layer - dre distal radular epithelium - dz degeneration zone - fe front edge - g granula - gol dictyosome - hd hemidesmosome - hl haemolymph - ie inferior epithelium - j jaw - ma tooth matrix - mc mineralizing cell - mem membranoblast - mfb microfibrills - mfl microfilaments - mgb multigranular body - mi mitochondria - mit mitosis - ml middle layer - mt microtubuli - mv microvilli - mw membrane whirl - n nucleus - nc necrotic cluster - nf nerve fibres - nsg neurosecretory granula - o odontophor - od odontoblast - odg odontoblast group - pod pre-odontoblast - rb residual body - rer rough endoplasmatic reticulum - rm radular membrane - rt radula teeth - sc supporting cell - se superior epithelium - sj septate junction - sro subradular organ - ss secretion substance - tf tonofilaments - tsm supramedian tensor muscle - tw terminal web - v vacuole - ves vesicle