Further investigations on the structure and function of cephalic organs of a nemertine Lineus ruber

The glandular cells of these paired neuroglandular organs pour their secretions (neutral mucopolysaccharide) into a cerebral (cephalic) canal connecting the organ lobule to the external medium. The secretions are then taken up by the 'vesicular cells' lining the blind end of the tube. They are transformed into acid -mucopolysaccharide in the vesicular cell and are discharged at the basal surface of the cell. The discharged materials are phagocytosed by cells of another type which resembles macrophages; these enter the blood lacuna by 'diapedesis' through the wall of the organ. The activity of the glandular cells and vesicular cells changes coincidentally with light and with salt concentration in the external medium. The function of the organ is discussed.     

Special cutaneous receptor organs of fish. 3. The ampullary organs of Eigenmannia

Ampullary receptor organs of the South American weakly electric gymnotid fish Eigenmannia virescens consist of a pore at the surface of the skin, a canal through the epidermis, and the expanded basal end of the canal in the corium. The cavity of the organ contains a jelly that is filled with fine fibers. The canal wall consists of three to six layers of flattened cells that appear to be derived from the adjacent skin. Along the lumen of the organ the cells are joined by tight junctions. Usually there are four spherical receptor cells in the base of the organ. They are innervated by single neural terminals. These organs are compared to tuberous receptor organs found in the same species, and the functional significance of the fine structure observed in these cells is discussed.     

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     

Chemoreceptor sensillum structure in Limulus

The chemoreceptors of Limulus polyphemus (L.) are polyneuronal sensilla found in the spines of the coxal gnathobases of each walking leg, the spines of the chilarial appendages, and the chelae of all the limbs. Each sensillum contains 6–15 bipolar sensory cells that share a single pore in the cuticle. The dendrites of the sensory cells of each sensillum course to the cuticle together. These attenuate sharply and enter a canal in the cuticle as a very narrow terminal thread. The dendrites retain their identity in the thread, but with the light microscope, they are usually not visible individually. Each thread, consisting of 6–15 dendrites, is accompanied to the cuticular surface by a cuticular tubule found within the canal. The chemoreceptor sensilla of the gnathobase, chilarium, and chela, the temperature organs of Patten, and the flabellar receptor organs all have the same basic organization. In general this is the same structural plan shown by chemoreceptors of other arthropods. Several different mechanisms of peripheral physiological interaction among receptor cells are possible with a sensillum organization like that described here for Limulus.     

A comparative ultrastructural investigation of the cephalic sensory organs in Opisthobranchia (Mollusca, Gastropoda)

Cephalic sensory organs (CSOs) are specialised structures in the head region of adult Opisthobranchia involved in perception of different stimuli. The gross morphology of these organs differs considerably among taxa. The current study aims at describing the cellular morphology of the CSOs in order to reveal cellular patterns, especially of sensory epithelia, common for opisthobranchs. Transmission electron microscopy was used to characterise the fine structure of the organs and to compare the CSOs of four different opisthobranch species. The cellular composition of the sensory system is conserved among taxa. The epidermal cells in sensory regions are always columnar and ciliated cells are frequently apparent. The sensory cells are primary receptors arranged in subepidermal cell clusters. They extend dendrites which penetrate the epithelium and reach the surface. Some of the dendrites bear cilia, whereas others only build a small protuberance. Processing of sensory information takes place in the peripheral glomeruli of all species. Moreover, few taxa possess additional peripheral ganglia at the base of their CSOs. The results of the present study might support other investigations indicating that the posterior CSOs are primarily involved in distance chemoreception, whereas the anterior CSOs might be used for contact chemoreception and mechanoreception.     

The hair-peg organs of the shore crab,Carcinus maenas (Crustacea,Decapoda): Ultrastructure and functional properties of sensilla sensitive to changes in seawater concentration

Summary The hair-peg organs of the shore crab, Carcinus maenas, are modified hair-sensilla. A small hair shaft (peg) is surrounded by a tuft of solid cuticular bristles (hairs). Each hair-peg organ is innervated by 6 sensory neurons, 2 of which have scolopidial (type-I) dendrites. The outer segments of all dendrites pass through a cuticular canal extending to the articulated hair base in which the 2 type-I dendrites terminate. The other 4 (type-II) dendrites reach the clavate tip of the hair shaft and have access to a terminal pore and a large sickle-shaped aperture. Three inner and 8–12 outer enveloping cells belong to a hair-peg organ. The innermost enveloping cell contains a scolopale, which has desmosomal connections to the ciliary rootlets of the type-I dendrites. An inner and an outer sensillum lymph space are present. The ultrastructural features of the dendrites and the cuticular apparatus indicate that the hair-peg organs are bimodal sensilla, comprising 2 mechano- and 4 chemosensitive sensory neurons. Extracellular recordings from the leg nerve indicate that the chemosensitive neurons of the hair-peg organs respond to changes in seawater concentration in the physiological range of Carcinus maenas.Supported by the Deutsche Forschungsgemeinschaft (SFB 45/A1; W. Gnatzy)     

Gustatory sense organs in the food canal of aphids

A sensory structure in the anterior region of the food canal of two species of aphid has been examined by light and electron microscopy. The dorsal wall is innervated by a total of 60 neurones which terminate, in groups, at 14 porous papillae on the cuticle. Paired papillae have also been detected in the ventral wall of this region. The fine structure of individual neurones and their grouping around papillae indicates a chemosensory function. The examination of moulting aphids shows that the distal portions of dendrites are shed with the exuviae.     

Ultrastructural investigation of the vesicular glands in Scutigera coleoptrata (Chilopoda,Notostigmophora)

In the notostigmophoran centipedes, two pairs of vesicular glands have evolved. These paired glands are situated in the first and second trunk segment and open via cuticular ducts in the upper part of the particular pleura. The vesicular glands of Scutigera coleoptrata were investigated using light and, for the first time, electron microscopical methods. The glands consist of wide sac‐like cavities that often appear vesicular. The epithelia of both glands are identically structured and consist of numerous glandular units. Each of these units consists of four different cells: a single secretory cell, a small intermediary cell, and one proximal and one distal canal cell. The intermediary cell forms a conducting canal and connects the secretory cell with the canal cells. Proximally, the intermediary cell bears microvilli, whereas the distal part is covered with a distinct cuticle. The cuticle is a continuation of the cuticle of the canal cells. This investigation shows that the ultrastructure of glandular units of the vesicular glands is comparable to that of the glandular units of other epidermal glands in Chilopoda and Diplopoda, although the glands look completely different in the light microscope. Thus, it is likely that the vesicular glands and epidermal glands share the same ground pattern. With regard to specific differences in the cuticular lining of the intermediary cells, a common origin of epidermal glands in Myriapoda and Hexapoda is not supported. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

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.     

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.     

A functional unit consisting of an eversible gland with neurosecretory innervation and a proprioreceptor derived from a complex sensillum in an insect

Summary The eversible sac of the antennal tip in Hypogastrura socialis (Collembola) has been reconstructed from serial ultrathin sections. The organ contains 3 specialized epidermal glandular cells and the dendrites of 2 sensory cells encapsulated by an enveloping cell. The following features of the system are particularly remarkable and have been analyzed in detail: (1) a neurosecretory innervation of the glandular cells, (2) the structure of the dendritic outer segments within the sac, and (3) the structure of the complex sensillum, from which these dendrites may be derived. The system may be thought of as providing an example of phylogenetic transformation of an exteroceptor into a mechanoreceptive proprioceptor. A functional model is proposed which involves control of the mechanism of evagination as well as of the secretory discharge.     

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 nuchal organs in the polychaete Platynereis dumerilii (Annelida,Nereididae)

Abstract. We examined the nuchal organs of adults of the nereidid polychaete Platynereis dumerilii by means of scanning and transmission electron microscopy. The most prominent features of the nuchal organs are paired ciliary bands located dorsolaterally at the posterior margin of the prostomium. They are composed of primary sensory cells and multiciliated supporting cells, both covered by a thin cuticle. The supporting cells have motile cilia that penetrate the cuticle and are responsible for the movement of water. Subapically, they have a narrowed neck region; the spaces between the neck regions of these supporting cells comprise the olfactory chamber. The dendrites of the sensory cells give rise to a single modified cilium that crosses the olfactory chamber; numerous thin microvillus-like processes, presumably extending from the sensory cells, also traverse the olfactory chamber. At the periphery of the ciliated epithelium runs a large nervous process between the ciliated supporting cells. It consists of smaller bundles of sensory dendrites that unite to form the nuchal nerve, which leaves the ciliated epithelium basally and runs toward the posterior part of the brain, where the perikarya of the sensory cells are located in clusters. The ciliated epithelium of the nuchal organs is surrounded by non-ciliated, peripheral epidermal cells. Those immediately adjacent to the ciliated supporting cells have a granular cuticle; those further away have a smooth cuticle. The nuchal organs of epitokous individuals of P. dumerilii are similar to those described previously in other species of polychaetes and are a useful model for understanding the development of nuchal organs in polychaetes.     

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.     

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.     

Vesicular structures in the dendrites of an insect olfactory receptor

The receptor dendrites of the antennal plaque organs of two species of fulgoroid insects contain 600 Å vesicles. These are concentrated in the distal vesicular region but also occur in the extreme tips of fine dendritic branches at the olfactory surface. At these points pore filaments run from the cuticular pores and are closely apposed to the dendritic membrane. Similar vesicles are found associated with Golgi bodies in the cell body and are also found more distally in multivesicular bodies. The multivesicular bodies appear to rupture and release vesicles. Vesicles also occur in the core of the dendrite cilium and it is assumed that they pass through the core and accumulate in the vesicular region. Similar vesicles appear to be a common feature in many, if not all, insect olfactory receptors.     

Fine structure of the sensory organs of Drosophila melanogaster Meigen larva (Diptera : Drosophilidae)

Fine structure of the prominent external and internal sensory organs of Drosophila melaogaster Meigan (Diptera : Drosophilidae) larva was determined by transmission electron microscopy (TEM). The external sensory organs, namely, antennal, maxillary, ventral and labial organs, dorsal pits, sensory cones on the 8th and 9th abdominal segments, and the sensory hairs on the body of the larva, were studied. A new knob in pit (KIP) sensillum innervated by 3 dendrites was found on the dorsolateral surface on either side. Four tufts of hairs at the posterior end of each great lateral tracheal trunk were found to be sensory with a dendrite at the base of each tuft.Internally in the pharynx of the larva, 3 groups of symmetrically located sensory organs, namely, the anteroventral, dorsal, and posteroventral groups were found. In all they contain 10 pairs of sensilla. Amongst them, 2 pairs of sensilla were found to be of the compound type; one having 9 dendrites arranged in 3 groups of 4, 3 and 2; while the other had 6 dendrites grouped as 2 and 4. In addition, 2 groups of sensilla were found on the internal dorsal fold on either side.Similarities were observed in the dendritic organisation of sensilla in the internal mouthparts of the Drosophila larva and the adult. Unlike nerves of the adult, the larval nerves connected with the dorsal and maxillary organs have a significantly thick layer of glial cells ensheathing the bundle of axons.     

Ultrastructure of the cerebral glands in Scolopendra cingulata Latr., Cryptops savignyi leach and C. hortensis Leach (Myriapoda : Scolopendromorpha)

The fine structure of the cerebral glands (cephalic neurohaemal organs) was investigated in the scolopendromorphs, Scolopendra sp. and Cryptops sp. (Myriapoda Scolopendromorpha). As in other Chilopoda, these organs are composed of glandular cells and axons, originating from the brain. Differences between the 2 genera studied concern: (i) The general structure (axons and glandular cells are mixed in Scolopendra and separated in Cryptops). (ii) The different cell types (I type of glial cell and 2 of glandular cells in Scolopendra while respectively 2 and 3 are persent in Cryptops. Fruthermore, this last genus has a less-developed Golgi apparatus and shows in one of the glandular cell types an intranuclear vacuole, sometimes related to a dilated perinuclear space. Physiological significance of such a structure remains unknown). (iii) The number of axonal types (less numerous in Scolopendra than in Cryptops). The 2 genera show little exocytosis profiles, both in axons and in glandular cells.     

Digitiform Sensilla on the Maxillar Palp of Coleoptera

The fine structure of the digitiform sensilla on the distal segment of the maxillar palps of Tenebrio and Dermestes is described. Each sensillum is associated with a single sensory cell and three enveloping cells, which enclose two receptor lymph cavities. The inner receptor lymph cavity of both species shows a different structural feature. Branches of the outer dendritic segments, which contain numerous microtubules, run to the tip of the hairshaft. A dendritic sheath extends to the apex of the peg. The hairshaft possesses a second canal, which is free of dendrites. The poreless hairshaft is inserted in a cuticular canal; the longer distal part of the shaft is positioned in a narrow superficial groove. The digitiform sensilla do not show the typical structures of mechanosensitive sensilla. The absence of pores in the setal wall does not point to a function as olfactory or gustatory hairs. The presumed function of the sensilla is discussed in relation to thermo-, hygro- and CO₂-receptors.