Ultrastructural investigation of the nuchal organs ofPygospio elegans (Polychaeta). I. Larval nuchal organs

The structural differentiation of the nuchal organs during the post-embryonic development ofPygospio elegans is described. The sensory organs are composed of two cell types: ciliated cells and bipolar primary sensory cells, constituting the nuchal ganglion, which is associated with both the sensory epithelium and the brain. Since the sensory neurons are largely integrated into posterolateral parts of the cerebral ganglion, the nuchal organs are primary presegmental structures. The microvilli of the ciliated cells form a cover over the cuticle with a presumed protective function. An extracellular space extends between cuticle and sensory epithelium. The distal dendrites of the sensory cells terminate in sensory bulbs, bearing one modified sensory cilium each that projects into the olfactory chamber, embedded within the secretion of the ciliated cells. During development, the nuchal organs increase in size. This is accompanied by a shift in position, an expansion of the sensory area, and secretory activity of the ciliated cells. The nuchal ganglion differentiates into three nuchal centres forming three distinct sensory areas around the ciliated region. Each nuchal complex reveals two short nuchal nerves comprising the sensory axons, which enter the posterior circumesophageal connective. The sensory cells lying in the brain exhibit neurosecretory activity; the sensory cilia enlarge their surface area by dilating and branching. Nuchal organs accomplish the basic structural adaptions of chemoreceptors and show structural analogies to arthropod olfactory sensilla; thus, there is every reason to suppose chemoreceptor function.     

Ultrastructure of the nuchal organ in the interstitial polychaete Stygocapitella subterranea (Parergodrilidae)

The nuchal organs of Stygocapitella subterranea are paired narrow pits. They are lined by unciliated cells at the opening and by ciliated cells at the basal parts. The primary sensory cells (6–8) are arranged in a single patch at the bottom of the nuchal pit. The nuclei of the sensory cells are located in the posterior portion of the brain. Their dendrites form the nuchal nerve which is sheathed by the ciliated cells. Each sensory cell bears up to 4 modified sensory cilia and several microvilli extending into the olfactory chamber. The sensory cilia show various patterns of axonemal organization and have no rootlets. The olfactory chamber is covered by a cuticular matrix. Another primary sensory cell lies at the opening of the nuchal pit. It bears cilia which penetrate the cuticle but are enveloped by the epicuticle. Retractor muscles insert caudally on the organ. The nuchal organ of S. subterranea shows similarities to those of opheliids but exhibits several features not to be found in other nuchal organs.     

Comparative ultrastructure of juvenile and adult nuchal organs of an annelid (Polychaeta: Opheliidae)

Opheliid nuchal organs are composed of ciliated cells, retractor muscles, and sensory cells. The perikarya of sensory cells are located in the posterior portion of the brain, and their distal processes extend along the body wall, as the nuchal nerve, and terminate just anterior to the ciliated region. The nuchal nerve of the juvenile is composed of 30–35 dendrites; the adult nuchal nerve has 35–40 dendrites. The ends of the sensory dendrites form sensory bulbs which are clustered around the olfactory chamber, and each bulb bears a modified cilium. Sensory cilia lose their axonemes and extend as microvillous-like structures into the olfactory chamber. Supportive cells delineate approximately the posterior and dorsal portions of the chamber with sensory bulbs forming the remaining ventral and anterior portions. On the lateral aspect of the chamber, cuticular matrix extends into it, and in this area supportive cells bear microvilli which extend into the matrix. The adult nuchal organ is larger than that of the juvenile, and the sensory portion of the olfactory chamber wall is expanded. Expansion of the sensory area is apparently the result of size increase in sensory bulbs and by intrusion of supportive cells between sensory bulbs.     

Ultrastructural investigations on the nuchal organ of the protandric polychaete,Ophryotrocha puerilis (Polychaeta,Dorvilleidae)

Summary The nuchal organs of the protandric hermaphrodite Ophryotrocha puerilis were studied by electron microscopy. Ophryotrocha puerilis is the first species hitherto described which possesses four instead of two nuchal organs. These sensory structures are located as ciliary pits at the posterior margin of the prostomium. Histologically, the nuchal organs are composed of supporting cells with long motile cilia and bipolar sensory cells, the perikarya of which form four distinct nuchal ganglia adjoining the brain. These structural components are concentrically arranged around the central sensory area. This area is covered by a modified cuticle, whereas the cuticle above the peripheral region of the sense organ exhibits the appearance typical for polychaetes. Two types of vesicular material are produced in the basal supporting cells, a dense-cored one within the central supporting cells only and a clear irregular-shaped one in all of these cells. The first type is considered to be responsible for the formation of the modified cuticle. The significance of these most probably long-distance chemoreceptory organs and their possible role in reproductive behaviour is discussed.     

Ultrastructure of nuchal organs in some marine polychaetes

Polychaetes normally possess one pair of nuchal organs at the posterior edge of the prostomium or peristomium. They have been regarded as chemosensory organs. The nuchal organs of four marine polychaete species with different habits were investigated by electron microscopy. Although the shapes of nuchal organs can vary greatly from simple ciliary bands (Scolelepis squamata, Spionidae) to retractile tongue-like, piston- or finger-shaped forms (Eteone longa, Anaitides mucosa, Phyllodocidae; Heteromastus filiformis, Capitellidae), the structural components, including the ciliated supporting cells, sensory cells, and nuchal epidermal cells, are essentially similar. The differences basically concern 1) the position of the sensory cells with relation to the ciliated supporting cells, 2) the location and structure of the nuchal nerve, and 3) the structure of the nuchal cuticle. The diverging nature of this modified cuticle is described and discussed in detail. Comparisons are made with the fine structure of nuchal organs of other polychaete species. Similarities of cellular components of nuchal organs are found not only in the four species studied here but also in all nuchal organs investigated so far. This is hypothesized to be due to the fact that the polychaete stem species already possessed nuchal organs with the respective cell types. Differences in the number and distribution of cellular components and in the overall shape of nuchal organs are thought to have evolved in correlation with the equipment of other cephalic appendages and with different habits and modes of nutrition.     

Ultrastructure of the extensively developed nuchal organs of Laonice bahusiensis (Annelida: Canalipalpata: Spionidae)

The nuchal organs of annelid Laonice bahusiensis (Spionidae) from northern Europe have been studied using scanning and transmission electron microscopy. L. bahusiensis is the first spionid species in which extensively developed, continuous nuchal organs are described. The nuchal organs of this genus are the longest known among polychaete annelids. They consist of paired double bands extending from the prostomium on a mid‐dorsal caruncle for about 24–30 setigers. Their microanatomy corresponds to the general structural plan of nuchal organs: there are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving‐stone‐like cover and innervated by one pair of longitudinally elongated nuchal nerves. These findings clearly favor the hypothesis that the paired, extensively developed ciliated structures found in some Spionidae are homologous with the prostomial nuchal organs characteristic of polychaete annelids. J. Morphol. 2010. © 2009 Wiley‐Liss, Inc.     

Ultrastructure of Nuchal Organs in Polychaetes (Annelida) — New Results and Review

Nuchal organs are epidermal sensory structures present in most polychaetes. They are situated at the posterior edge of the prostomium and may extend posteriorly onto the peristomium. Although there is considerable external variation, they all consist of ciliated supporting cells, bipolar primary sensory cells and retractor muscles. They are innervated directly from the brain by paired nerves. The sensory cells are usually monociliated; their sensory processes lie in subcuticular spaces, the olfactory chambers. Structural variability is to be observed in the location of the sensory cells, the course of the nuchal nerve, position of nuchal ganglia as well as in cytological features of sensory and supporting cells. These differences provide useful characters for phylogenetic considerations to establish supraspecific taxa within the phylogenetic system of the Annelida. Special emphasis is laid on the problem of whether the nuchal organs represent an autapomorphy of the Polychaeta or the Annelida and thus whether the lack of nuchal organs in Clitellata is primary or secondary. As is discussed, the probability of a loss of the nuchal organs in Clitellata is higher, which favours the second hypothesis: that nuchal organs are part of the ground pattern of the Annelida and very likely are an autapomorphy of this group.     

Ultrastructural investigation of the nuchal organs of Pygospio elegans (Polychaeta)

Summary The differentiation of the dorsal organs as well as the structure of the nuchal organs and their relation to the central nervous system in adult Pygospio elegans were studied by electron microscopy and compared to the nuchal organs of the larvae. The nuchal organs are represented by paired ciliary bands on the dorsal side of the first setiger, delimiting a median caruncle that is completely filled with epidermal and nervous tissue. They are composed of ciliated supporting cells and bipolar primary sensory cells constituting the nuchal ganglia, which are integrated into the brain. Microvillus-like processes of the ciliated cells give rise to a secondary covering layer over the sensory epithelium. The size of the nuchal organs is a sexually dimorphic feature.Dorsal organ formation is concomitant with the onset of sexual maturation in the male sex only. They appear as metameric ciliary bands on the dorsal side of the anterior body region and consist of ciliated cells accompanied by lateral accumulations of tubular gland cells. In the gametogenic segments they are structurally associated with the male genital pores and may be involved in reproduction. The results refute previous theories that dorsal organs are sensory and have a common origin to nuchal organs.Abbreviations ac anterior commissure of the brain - ace anterior circumesophageal connective - bb basal body - bl basal lamina - c cuticle - ca caruncle - cc ciliated cell - ci sensory cilium - co microvillar cover - d septate desmosome - db dorsal blood vessel - dn dorsal nerve cord - ea efferent axons - ec epidermal cell - eg elementary granules - g Golgi complex - i filamentous inclusion - lm longitudinal muscles - ly lysosome - mc motile cilia - mv microvillus - n neuron - ng nuchal ganglion - nn nuchal nerve - nu nucleus - oc olfactory chamber - pa palp - pc posterior commissure of the brain - pce posterior circumesophageal connective - rer rough endoplasmic reticulum - sI setiger I - sb sensory bulb - sc sensory cell - sd sensory dendrite - ser smooth endoplasmic reticulum - tf tonofilament bundle - v clear vesicles - za zonula adherens     

Ultrastructural investigations on the cephalic and metameric nuchal organs of Spio cf. filicornis (Polychaeta, Spionidae)

The nuchal organs of Spio cf. filicornis from northern Europe have been studied by scanning and transmission electron microscopy. Spio cf. filicornis is the first species in which metameric nuchal organs are described. The nuchal organs consist of a distinct cephalic nuchal complex followed by metameric structures for a variable number of chaetigers. Their microanatomy corresponds to the general structural plan of nuchal organs: these are ciliated supporting cells and bipolar sensory cells with sensory cilia traversing an olfactory chamber. The organs are overlaid by a secondary paving-stone-like cover and innervated by longitudinally elongated paired nuchal nerves. The findings clearly favour the hypothesis that the paired metameric ciliated structures found in some Spionidae are in fact homologous with the prostomial nuchal organs characteristic of Polychaeta.     

Olfactory metamorphosis in the coastal tailed frog Ascaphus truei (Amphibia,Anura, Leiopelmatidae)

The structure of the olfactory organ in larvae and adults of the basal anuran Ascaphus truei was examined using light micrography, electron micrography, and resin casts of the nasal cavity. The larval olfactory organ consists of nonsensory anterior and posterior nasal tubes connected to a large, main olfactory cavity containing olfactory epithelium; the vomeronasal organ is a ventrolateral diverticulum of this cavity. A small patch of olfactory epithelium (the “epithelial band”) also is present in the preoral buccal cavity, anterolateral to the choana. The main olfactory epithelium and epithelial band have both microvillar and ciliated receptor cells, and both microvillar and ciliated supporting cells. The epithelial band also contains secretory ciliated supporting cells. The vomeronasal epithelium contains only microvillar receptor cells. After metamorphosis, the adult olfactory organ is divided into the three typical anuran olfactory chambers: the principal, middle, and inferior cavities. The anterior part of the principal cavity contains a “larval type” epithelium that has both microvillar and ciliated receptor cells and both microvillar and ciliated supporting cells, whereas the posterior part is lined with an “adult‐type” epithelium that has only ciliated receptor cells and microvillar supporting cells. The middle cavity is nonsensory. The vomeronasal epithelium of the inferior cavity resembles that of larvae but is distinguished by a novel type of microvillar cell. The presence of two distinct types of olfactory epithelium in the principal cavity of adult A. truei is unique among previously described anuran olfactory organs. A comparative review suggests that the anterior olfactory epithelium is homologous with the “recessus olfactorius” of other anurans and with the accessory nasal cavity of pipids and functions to detect water‐borne odorants. J. Morphol. 2011. © 2011 Wiley Periodicals, Inc.     

Histology and ultrastructure of the olfactory organ of the freshwater pulmonate Helisoma trivolvis

Summary The olfactory organ of Helisoma trivolvis is located on the surface of the body at the base of the cephalic tentacles. An evagination of skin, the olfactory plica, at the base of the tentacle extends over the olfactory organ dorsally. The epithelium of the olfactory organs contains unspecialized epithelial cells, ciliated epithelial cells, basal cells, mucous secretory cells, and sensory dendrites. The surface of the epithelium has a complex brush border of thick plasmatic processes, which branch to form several terminal microvillar twigs. Long slender cytoplasmic processes form a dense spongy layer among the plasmatic processes beneath the level of the terminal twigs. Bipolar primary sensory neurons clustered beneath the epithelium of the olfactory organ send dendrites through the epithelium to the free surface. Some sensory endings have a few short cilia, but most bear only microvilli. Cilia of sensory endings and epithelial cells extend beyond the brush border of the epithelium. Small axons arise from the perikarya of the sensory neurons and enter a branch of the olfactory nerve. HRP tracing indicates that the axons pass to the cerebral ganglion without interruption. Histochemical tests indicate that the sensory neurons are neither aminergic nor cholinergic.     

Ultrastructure of cell types of the olfactory epithelium in a catfish,Heteropneustesfossilis (Bloch)

Transmission electron microscopical study of olfactory epithelium of a mud-dwelling catfish,Heteropneustes fossilis (Bloch) shows receptor, supporting, goblet and basal cells. The receptor cells are of ciliated and microvillous type. Both ciliated and microvillous receptor cells are provided with olfactory knob. The dendrite of all the receptor cells bears many longitudinally arranged microtubules. Occurrence of the rod cell and its function is quite debatable. Specialized juctional complexes between the receptor and adjacent cells are clearly noted. The supporting cells are both ciliated and nonciliated. The ciliated supporting cells are responsible for water ventilation in the olfactory chamber as well as in the inter-lamellar spaces. This facilitates better perception of odours by the receptor cells. In addition to providing mechanical support to other cells, the nonciliated supporting cells also have a secretory function which is evident from the present study. The different stages of maturity of goblet cells are well documented. The presence of white cells in the olfactory epithelium is a very rare finding.     

Ultrastructure of the nuchal organ and cerebral organ in Onchnesoma squamatum (Sipuncula, Phascolionidae)

 The ultrastructure of the nuchal organ and cerebral organ is described for the first time in a species of the Sipuncula, Onchnesoma squamatum. The nuchal organ is an unpaired structure lying outside and dorsal to the tentacular crown; furrows give the organ a paired appearance. The cerebral organ is an unciliated pad anterior to the nuchal organ. The nuchal organ consists of ciliated supporting cells, non-ciliated supporting cells and bipolar primary sensory cells. The cerebral organ is composed of unciliated supporting cells and numerous bipolar sensory cells. This clearly favours the hypothesis that this structure has a sensory function in adults rather than being a vestige of a larval organ. The sensory cells are similar in both organs and exhibit features indicative of chemoreception. Since the density of the sensory cells is low in the nuchal organ, an exclusively sensory function is questioned. There is some evidence that the two organs represent a functional unit. The present findings do not support the view that the nuchal organs of Sipuncula and ”Polychaeta” are homologous, but instead suggest that they are convergent structures. Accepted: 18 September 1996     

Anatomy,histology, and histochemistry of the olfactory organ of the Korean shuttles mudskipper Periophthalmus modestus

The Korean shuttles mudskipper Periophthalmus modestus has paired olfactory organs on its snout, consisting of anterior and posterior nostrils, a single olfactory canal with sensory and nonsensory epithelia, and a single accessory nasal sac. Its sensory epithelium consists of numerous islets forming a pseudostratified layer and contains various cells: olfactory receptor neurons, supporting cells, basal cells, lymphatic cells (LCs), and axon bundles. The sensory epithelium is a stratified squamous layer comprising stratified epithelial cells, mucous cells (MCs) with glycogen, flattened cells (FCs), LCs, and unidentified cells. Specific structures are as follows: (a) a tubular anterior nostril projecting outward, (b) a slit posterior nostril, (c) an elongated olfactory canal, (d) an ethmoidal accessory nasal sac, (e) axon bundles found only in the basal layer of the sensory epithelium, (f) FCs only at the top of the nonsensory epithelium, and (g) glycogen-containing MCs. Such structures seem to be unique in that they have not been observed in most teleost fishes spending their whole life in water.     

Sense organs in polychaetes (Annelida)

Polychaetes possess a wide range of sensory structures. These form sense organs of several kinds, including the appendages of the head region (palps, antennae, tentacular cirri), the appendages of the trunk region and pygidium (parapodial and pygidial cirri), the nuchal organs, the dorsal organs, the lateral organs, the eyes, the photoreceptor-like sense organs, the statocysts, various kinds of pharyngeal papillae as well as structurally peculiar sensory organs of still unknown function and the apical organs of trochophore larvae. Moreover, isolated or clustered sensory cells not obviously associated with other cell types are distributed all over the body. Whereas nuchal organs are typical for polychaetes and are lacking only in a few species, all other kinds of sensory organs are restricted to certain groups of taxa or species. Some have only been described in single species till now. Sensory cells are generally bipolar sensory cells and their cell bodies are either located peripherally within the epidermis or within the central nervous system. These sensory cells are usually ciliated and different types can be disinguished. Structure, function and phylogenetic importance of the sensory structures observed in polychaetes so far are reviewed. For evaluation of the relationships of the higher taxa in Annelida palps, nuchal organs and pigmented ocelli appear to be of special importance.     

Surface ultrastructure of the olfactory rosette of an air-breathing catfish,Heteropneustes fossilis (Bloch)

The surface architecture of the olfactory rosette ofHeteropneustes fossilis (Bloch) has been studied by scanning electron microscopy. The olfactory rosette is an oval structure composed of a number of lamellae arranged pinnately on a median raphe. The raphe is invested with epithelial cells and pits which represent goblet cell openings. On the basis of cellular characteristics and their distribution the lateral surface of each olfactory lamella is identified as sensory, ciliated non-sensory and non-ciliated non-sensory epithelium. The sensory epithelium is provided with receptor and supporting cells. The ciliated non-sensory epithelium is covered with dense cilia obscuring the presence of other cell types. The non-ciliated non-sensory epithelium is with many polygonal areas containing cells.     

Lateral organs in sedentary polychaetes (Annelida) – Ultrastructure and phylogenetic significance of an insufficiently known sense organ

Lateral organs are sense organs visible as densely ciliated pits or papillae between the noto‐ and the neuropodia in certain taxa of sedentary polychaetes. Ultrastructural studies in about 10 species of the following taxa Maldanidae, Opheliidae, Orbiniidae, Paraonidae, Magelonidae, Spionidae, Poecilochaetidae and Terebellidae have been designed to evaluate whether these organs are homologous among polychaetes. In spite of great external diversity, the investigations revealed an overall ultrastructural similarity. Differences between species investigated mainly concern the size of the organs as well as the number and arrangement of cells. The organs comprise supportive cells and uniciliated penetrative sensory cells. Their dendrites are closely arranged and thus their cilia may resemble multiciliated cells. There are two types of sensory cells: one type possesses no or mainly thin microvilli of which usually only a few reach the cuticular surface, and in the other type the cilium is consistently surrounded by 10 strong microvilli, which form a pore‐like opening in the cuticle. Further differences occur in the structure of the rootlet system. Basally, a retractor muscle attaches to the organ. The systematic significance of these organs within Annelida is discussed with respect to the conflicting phylogenetic hypotheses explaining the relationships of annelid taxa.     

黑斑蛙嗅器和犁鼻器的组化及超微结构特征

嗅感受器主要感知外界环境中化学信号分子.本文采用银染、NADPH-组化染色和电镜技术来观察黑斑侧褶蛙(Petophylax nigromaculatus)的嗅器和犁鼻器的功能差异及细胞组成.银染法可对嗅上皮和犁鼻上皮的细胞进行分类及区分.其中,支持细胞胞核深染成黑色,嗅细胞胞核银染为花斑状.细胞计数显示,犁鼻上皮的嗅神经细胞含量百分比显著高于嗅上皮.组化结果显示,黑斑侧褶蛙嗅上皮和犁鼻上皮对NADPH-d表达模式差异显著,前者表达明显高于后者.电镜结果显示,黑斑侧褶蛙嗅上皮和犁鼻上皮的支持细胞由两种类型的细胞组成,分别为纤毛型和颗粒型支持细胞.     

The ultrastructural organization of olfactory epithelium of two species of gadoid fish

The untrastructural organization of the olfactory epithelium of the cod Gadus morhua (L.) and the haddock Melanogrammus aeglefinus (L.) was studied using both transmission and scanning electron microscopy. The olfactory rosette was found to exhibit regional differences; the faces of the olfactory lamella were composed of sensory epithelium, the edges were non-sensory. The cellular organization of the olfactory epithelium was determined and consisted of bi-polar sensory neurones, supporting cells, mucous cells and basal cells. The ultrastructure of the sensory cells was consistent, having an elongate cell, the free surface of which terminated in an olfactory vesicle from which arose either four olfactory cilia or numerous microvilli. Ciliary aggregations have been found in the two species of gadoid fish studied; it is suggested that these structures aid in the separation and in the circulation of fluid between the lamellae. The surface structure of the supporting cells was found to be of two types: either ciliated or ridged; the former presenting distinct ciliated tufts, the latter showing definite, but unorganized, ridges over the epithelium surface.     

Ecostructural studies on olfactory organ in young and adult sea trout (Osteichthyes,Salmonidae)

Summary The development of cells of the olfactory organ in young (parr and smolt) and adult Baltic sea trout is investigated from an ecological point of view. A theory on cell dynamics is presented. Blastema cells are basal cells which divide into goblet cells, primary receptors and primary supporting cells. The latter cell type reminds one of undifferentiated fibroblasts. In the indifferent epithelium the primary supporting cells then differentiate into ciliated nonsensory cells, whereas in the sensory epithelium they form ciliated and nonciliated supporting cells. The primary receptors develop into spindle receptors, and these into rod receptors. This dimorphism of supporting cells and receptors and the grouping of them is discussed. Indications on aging of the epithelium are presented. The epithelial cells are arranged in zones. Mainly in the basal zone there is a supply of lymphoid wandering cells and macrophages some of which move into other zones and phagocytize dead or degenerating cells. Dead cells are also extruded into the olfactory chamber. This occurs especially during the initial secondary folding of the primary olfactory laminae. Different aspects on ecological adaptation of cells and structures are presented. Baltic sea trout have a dynamic cell population and secretion adapted for life both in river and sea.