Ultrastructure of tarsal sensilla and other integument structures of two Pseudocellus species (Ricinulei, Arachnida)

Ricinuleids are one of the least investigated groups of Arachnida. In particular, knowledge of their ultrastructure is poor. Observations of the distal tarsomeres of ricinuleids show differences in their shape and equipment of surface structures. Legs I and II are used by the Ricinulei to explore their surroundings with tentative movements. The tarsomeres of these legs show similarities in shape and surface structures that distinguish them from those of legs III and IV. In this study, 11 different structures of the tarsomere surfaces of two cave-dwelling species, Pseudocellus pearsei and P. boneti from México, were investigated for the first time with scanning and transmission electron microscopy and discussed regarding their possible function: 1) a single treelike ramifying seta resembles a no pore single-walled (np-sw) sensillum; 2) setae occurring in a small number and possessing a bipartite shaft represent terminal pore single-walled (tp-sw) sensilla. The surface of the proximal half of the shaft shows small branches. The distal half has a smooth surface; 3) long setae with conspicuous longitudinal lamellae show characteristics of chemoreceptive wall pore single-walled (wp-sw) sensilla; 4) frequent small wp-sw sensilla with flat and irregular lamellae; 5) very short wp-sw sensilla occurring solitary or in groups; 6) a few short setae with smooth surface correspond to wp-sw sensilla; 7) a single short clubbed seta articulating in a flat pit is considered to be a np-sw sensillum; 8) common long setae with a pointed tip show characteristics of mechanoreceptive np-sw sensilla; 9) ventral setae with adhesive and mechanosensory function are accompanied by multicellular "class III" glands; 10) slit organs with mechanoreceptive function; and 11) dome-like tubercles with no indication of sensorial function. Several of these sensilla form a sensory field on the dorsofrontal surface which is particularly pronounced on the distal tarsomeres of legs I and II.     

The sensory equipment of a spider – A morphological survey of different types of sensillum in both sexes of Argiope bruennichi (Araneae,Araneidae)

Spiders show a wide range of sensory capabilities as evidenced by behavioural observations. Accordingly, spiders possess diverse sensory structures like mechano-, hygro-, thermo- or chemoreceptive sensilla. As to chemoreceptive structures, only trichoid tip-pore sensilla were found so far that were tested for gustation. That spiders are also able to receive airborne signals is corroborated by numerous behavioural experiments but the responsible structures have not been determined yet. Here, we provide sensilla distribution maps of pedipalps and walking legs of both sexes of the wasp spider Argiope bruennichi whose biology and mating system is well explored. By means of scanning electron microscopy, we scrutinized whether there is in fact only one type of trichoid pore sensillum and if so, if there are deviations in the outer structure of the tip-pore sensilla depending on their position on the body. We also describe the external structure and distribution of slit sense organs, trichobothria and tarsal organs. Our study shows that all four sensillum types occur on pedipalps and walking legs of both sexes. As to chemosensory organs, only tip-pore sensilla were found, suggesting that this sensillum type is used for both gustation and olfaction. The highest numbers of tip-pore sensilla were observed on metatarsi and tarsi of the first two walking legs. Mechanosensitive slit sense organs occur as single slit sensilla in rows along all podomers or as lyriform organs next to the joints. The mechanosensitive trichobothria occur on the basal part of tibiae and metatarsi. Tarsal organs occur on the dorsal side of all tarsi and the male cymbium. The distribution maps of the sensilla are the starting point for further exploration of internal, morphological differences of the sensilla from different regions on the body. Cryptic anatomical differences might be linked to functional differences that can be explored in combination with electrophysiological analyses. Consequently, the maps will help to elucidate the sensory world of spiders.     

Fine-structural characteristics of the antennal sensilla of Agrotis segetum (Insecta: Lepidoptera)

Summary The turnip moth Agrotis segetum possesses seven different types of sensilla: four single-walled (SW), one double-walled (DW), one terminal-pore (TP), and one poreless sensilla (NP).The SW 1 and SW 2 sensilla have the same external appearance, being long and slender, but differ in the branching pattern of the sensory processes: unbranched and branched in SW 1 and SW 2, respectively. The SW 3 sensilla are shorter, sickle-shaped, and contain a large number of branches from the sensory processes. These three sensillar types are innervated by 2–3 sensory cells. The SW 4 sensilla are raisin-shaped and possess three profusely branched sensory processes. The DW sensilla are short and have apical slit-like pores. This sensillar type has 5–6 sensory processes. The TP sensilla possess five sensory processes, one of them terminates basally in a tubular body, the others in the apical part of the long cuticular bristle. The NP sensilla are stout and have apical conelike structures. Two of the sensory processes terminate in the apical part, the third proximally. The third sensory process has a lamellar pattern. The fine structure indicates the following functions: SW and DW sensilla: chemoreception; TP sensillum: chemoreception and mechanoreception; NP sensillum: thermoreception and hygroreception.Supported by joint grants from the Swedish Council for Planning and Coordination of Research, the Swedish Natural Science Research Council, and the Swedish Council for Forestry and Agricultural Research     

Ultrastructure of chemosensory tarsal tip-pore sensilla of Argiope spp. Audouin, 1826 (Chelicerata: Araneae: Araneidae)

While chemical communication has been investigated intensively in vertebrates and insects, relatively little is known about the sensory world of spiders despite the fact that chemical cues play a key role in natural and sexual selection in this group. In insects, olfaction is performed with wall–pore and gustation with tip-pore sensilla. Since spiders possess tip-pore sensilla only, it is unclear how they accomplish olfaction. We scrutinized the ultrastructure of the trichoid tip-pore sensilla of the orb weaving spider Argiope bruennichi—a common Palearctic species the males of which are known to be attracted by female sex pheromone. We also investigated the congener Argiope blanda. We examined whether the tip-pore sensilla differ in ultrastructure depending on sex and their position on the tarsi of walking legs of which only the distal parts are in contact with the substrate. We hypothesized as yet undetected differences in ultrastructure that suggest gustatory versus olfactory functions. All tarsal tip-pore sensilla of both species exhibit characters typical of contact-chemoreceptors, such as (a) the presence of a pore at the tip of the sensillum shaft, (b) 2–22 uniciliated chemoreceptive cells with elongated and unbranched dendrites reaching up to the tip-pore, (c) two integrated mechanoreceptive cells with short dendrites and large tubular bodies attached to the sensillum shaft's base, and (d) a socket structure with suspension fibres that render the sensillum shaft flexible. The newly found third mechanoreceptive cell attached to the proximal end of the peridendritic shaft cylinder by a small tubular body was likely overlooked in previous studies. The organization of tarsal tip-pore sensilla did not differ depending on the position on the tarsus nor between the sexes. As no wall-pore sensilla were detected, we discuss the probability that a single type of sensillum performs both gustation and olfaction in spiders.     

Sensilla on the palps and legs of the adult soft tick argas persicus oken (IXODOIDEA: ARGASIDAE) and their projections to the central nervous system

The sensory structures present on the palps and legs of adult Argas persicus Oken (Ixodoidea: Argasidae) were studied by light, scanning and transmission electron microscopy. The number, distribution, surface morphology and the fine structure of the prominent sensilla present on these appendages were determined. The palps have 2 morphologically prominent types of sensilla: one with a grooved surface of the hair and the other having a non-grooved hair. The TEM distinguishes at least 4 prominent subtypes in grooved sensilla with single or double lumina and dendrites occupying the periphery of the central lumen or distributed all over the central lumen. Amongst the sensilla with non-grooved hair-shaft, a rare type of Olfactory Mechanoreceptive (OM) sensillum was found on the palps and the first legs of A. persicus. At the base of the hair-shaft, the OM sensillum has 2 mechanosensory dendrites. The hair-shaft of the sensillum has a porous cuticle, characteristic of an olfactory sensillum. The lumen of the hair-shaft is invested with branching dendrites from 3–8 neurons, which are surrounded by 4 sheath cells. The sensilla on the legs, including those present in the Hallers organ, are of at least 3 prominent categories. (i) Single wall with un-innervated hair-shaft. (ii) Single wall, multiporous sensillum with dendrites present in the hair shaft. (iii) Double walls with spoke channels and dendrites present in the central lumen. Sensory projections from the crown of sensilla located on the distal end of the palp extend to the palpal and suboesophageal (SOG) ganglia. Projections in the SOG extend further to the contralateral side. Sensilla in the Hallers organ project to the first pedal ganglion and to the anterodorsal region of supraoesophageal ganglion. As expected, the primary sensory projections from the sensilla of the other 3 legs extend to the respective pedal ganglia.     

Sensory Organs in Ips typographus (Insecta: Coleoptera) Fine structure of the sensilla of the maxillary and labial palps*

The labial and maxillary palps of the bark beetle, Ips typographus, possess a diversified array of sensilla. There are four types of sensilla possessing a single tubular body indicating a mechanoreceptive function. The variation of the associated cuticular structures of these sensillar types ranges from long bristles to cavities within the cuticle. Terminal pore sensilla with a supposed mechanosensory/gustatory function and single-walled presumably chemoreceptive sensilla are found on the apical part of the terminal palp segments. A poreless sensillar type is found on the lateral side of the terminal segment of the maxillary palp. The functional capabilities of this sensillar type are presently unknown.     

Sensing the Underground - Ultrastructure and Function of Sensory Organs in Root-Feeding Melolontha melolontha (Coleoptera: Scarabaeinae) Larvae

Introduction

Below ground orientation in insects relies mainly on olfaction and taste. The economic impact of plant root feeding scarab beetle larvae gave rise to numerous phylogenetic and ecological studies. Detailed knowledge of the sensory capacities of these larvae is nevertheless lacking. Here, we present an atlas of the sensory organs on larval head appendages of Melolontha melolontha. Our ultrastructural and electrophysiological investigations allow annotation of functions to various sensory structures.

Results

Three out of 17 ascertained sensillum types have olfactory, and 7 gustatory function. These sensillum types are unevenly distributed between antennae and palps. The most prominent chemosensory organs are antennal pore plates that in total are innervated by approximately one thousand olfactory sensory neurons grouped into functional units of three-to-four. In contrast, only two olfactory sensory neurons innervate one sensillum basiconicum on each of the palps. Gustatory sensilla chaetica dominate the apices of all head appendages, while only the palps bear thermo-/hygroreceptors. Electrophysiological responses to CO₂, an attractant for many root feeders, are exclusively observed in the antennae. Out of 54 relevant volatile compounds, various alcohols, acids, amines, esters, aldehydes, ketones and monoterpenes elicit responses in antennae and palps. All head appendages are characterized by distinct olfactory response profiles that are even enantiomer specific for some compounds.

Conclusions

Chemosensory capacities in M. melolontha larvae are as highly developed as in many adult insects. We interpret the functional sensory units underneath the antennal pore plates as cryptic sensilla placodea and suggest that these perceive a broad range of secondary plant metabolites together with CO₂. Responses to olfactory stimulation of the labial and maxillary palps indicate that typical contact chemo-sensilla have a dual gustatory and olfactory function.     

Sensory projections of identified coxal hair sensilla of the scorpionHeterometrus fulvipes (Scorpionidae)

The topography of long hair sensilla on the coxae of walking legs and pedipalps of the scorpionHeterometrus fulvipes is described. Identified long hair sensilla are cobalt filled, and central projections of sensory fibres are reported for the first time in the suboesophageal ganglion of this scorpion. The afferent fibres arising from each long hair sensilla segregate into ventral, dorsomedial and dorsal tracts upon their entry into the suboesophageal ganglion. These transverse tracts bifurcate towards the middle of the leg neuromere and form three ipsilateral, plurisegrnental, longitudinal sensory pathways. Filling a pair of bilaterally distributed long hair sensilla shows bilaterally arranged longitudinal afferent tracts interconnected by distinct transverse commissures. Similar patterns of sensory projections are observed when filling homologous hairs on other legs and pedipalps. Numerous fine collaterals arise from the longitudinal sensory trancts that subdivide and end in small blebs presumed to be presynaptic endings. The dorsal and dorsomedial longitudinal tracts and their respective commissures are in close association with the dendritic arborisations of pedipalpal and leg motor neurons, suggesting direct contact between them. The probable functions of these multisegmental hair afferent pathways are discussed.     

Female sex pheromone of a wandering spider (Cupiennius salei): identification and sensory reception

Females of the wandering spider Cupiennius salei attach a sex pheromone to their dragline. Males encountering the female dragline examine the silk thread with their pedipalps and then typically initiate reciprocal vibratory courtship with the sexual partner. The female pheromone was identified as (S)-1,1'-dimethyl citrate. The male pheromone receptive sensory cells are located in tip pore sensilla and respond to touching the sensillum tip with female silk or pieces of filter paper containing the synthetic pheromone.     

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.     

Antennal sensilla and their possible functions in the host-plant selection behaviour of Phenacoccus manihoti (Matile-Ferrero) (Homoptera : Pseudococcidae)

Nine different types of sensilla have been identified on the antenna of the cassava mealybug Phenacoccus manihoti (Homoptera : Pseudococcidae) with scanning and transmission electron microscopes. Trichoid sensilla, distributed on all segments of the antenna and innervated by a single mechanoreceptive dendrite, have the characteristics of exteroceptors. A campaniform sensillum located on the pedicel and one basiconic sensillum on the flagellum have the characteristics of proprioceptors. Coeloconic sensilla, located ventrally on the pedicel and flagellum, related to poreless sensilla with inflexible sockets, have the characteristics of thermo/hygroreceptors. Uniporous sensilla with a mechanoreceptive dendrite (smooth pegs P1 and P2, grooved pegs P3) and multiporous chemosensilla (grooved pegs P4 and P5), present on the tip of the flagellum, have, respectively, the characteristics of gustatory and olfactory receptors. The results of this study seem to suggest that the cassava mealybug has sensory equipment on its antennae that can detect, by olfaction and contact, chemicals released by the plant.     

Innervation of the cercal sensilla on the ovipositor of the Australian sheep blowfly (Lucilia cuprina)

ABSTRACT. The ovipositor of the female sheep blowfly, Lucilia cuprina (Wied.) (Diptera: Calliphoridae), has a complement of cercal sensilla that includes long, medium and short tactile hairs, two campaniform domes, four olfactory pegs, and ten double-channelled gustatory hairs. This sensory array is suited to assess oviposition site resources, prior to and during the laying of an egg batch.
The tactile hairs and campaniform sensilla are each innervated by a single, tubular body containing dendrite. The olfactory pegs are each innervated by a single, moderately branched dendrite, which gains access to the external environment via pores at the bottom of deep grooves in the peg wall. The gustatory hairs fall into two categories. Four hairs have a single, tubular body containing dendrite at their base, and four unbranched dendrites running up to the hair tip which has a terminal pore. Six of the taste hairs have no tubular body containing dendrite at the base, and three unbranched dendrites running up to a terminal pore.     

Spatially restricted expression of candidate taste receptors in the Drosophila gustatory system

BACKGROUND: Taste is an important sensory modality in most animals. In Drosophila, taste is perceived by gustatory neurons located in sensilla distributed on several different appendages throughout the body of the animal. Here we show that the gustatory receptors are encoded by a family of at least 54 genes (Gr genes), most of which are expressed exclusively in a small subset of taste sensilla located in narrowly defined regions of the fly's body. RESULTS: BLAST searches with the predicted amino acid sequences of 6 7-transmembrane-receptor genes of unknown function and 20 previously identified, putative gustatory receptor genes led to the identification of a large gene family comprising at least 54 genes. We investigated the expression of eight genes by using a Gal4 reporter gene assay and found that five of them were expressed in the gustatory system of the fly. Four genes were expressed in 1%-4% of taste sensilla, located in well-defined regions of the proboscis, the legs, or both. The fifth gene was expressed in about 20% of taste sensilla in all major gustatory organs, including the taste bristles on the anterior wing margin. Axon-tracing experiments demonstrated that neurons expressing a given Gr gene project their axons to a spatially restricted domain of the subesophageal ganglion in the fly brain. CONCLUSIONS: Our findings suggest that each taste sensillum represents a discrete, functional unit expressing at least one Gr receptor and that most Gr genes are expressed in spatially restricted domains of the gustatory system. These observations imply the potential for high taste discrimination of the Drosophila brain.     

The fine structure of the antennal sensilla of the pine saw flyNeodiprion sertifer (Insecta: Hymenoptera)

Summary There is a considerable sexual dimorphism as regards the development of the antenna inNeodiprion sertifer. In the male this is manifested in a larger antennal surface area and the possession of a great number of long single-walled sensilla (SW 1), which are located on the antennal branches. In the female the antennal branches are vestigial, and the single-walled sensilla, of a second type (SW 2), are shorter and present in lower numbers. Both sexes have in addition four types of sensilla in common:viz. a third type of single-walled sensilla (SW 3), terminal pore sensilla, double-walled sensilla, and poreless sensilla. These four sensillar types are characterized not only by their external appearance but also by their internal morphology. Especially the cells enveloping the sensory cells vary in number and appearance.Supported by joint grants from the Swedish Council for Planning and Coordination of Research, the Swedish Natural Science Research Council, and the Swedish Council for Forestry and Agricultural Research.     

Distribution of chemo- and mechanoreceptors on the antennae and maxillae of Busseola fusca larvae

The stem borer Busseola fusca (Fuller) (Lepidoptera: Noctuidae) is a major pest of maize, Zea mays L., and sorghum, Sorghum bicolor (L.) Moench (both Poaceae), in sub-Saharan Africa. Like in many other lepidopteran insects, the success of B. fusca in recognizing and colonizing a limited variety of plants is based on the interaction between its sensory systems and the physicochemical characteristics of its immediate environment. The sensilla on the maxillary galeae of B. fusca larvae are typical of Lepidoptera and comprise two uniporous styloconic sensilla, which are contact chemoreceptors, three basiconic sensilla, and two aporous sensilla chaetica. The maxillary palp is two-segmented and has eight small basiconic sensilla at the tip, which were also found to be gustatory. The antennae of B. fusca larvae are short and simple. The sensilla of the antenna are composed of two aporous sensilla chaetica, three multiporous cone-shaped basiconic sensilla, three small basiconic sensilla, and one aporous styloconic sensillum. The basiconic sensillum located on the third antennal segment displayed a contact chemoreception response. The other basiconic sensilla did not show any action potential activity in tip-recording tests. The significant and positive dose–response curve obtained for the antennal basiconic sensillum with sucrose indicated for the first time the presence of gustatory chemoreceptors on the antennae of a lepidopteran larva.     

The organization of the chemosensory system in Drosophila melanogaster: a rewiew

This review surveys the organization of the olfactory and gustatory systems in the imago and in the larva of Drosophila melanogaster, both at the sensory and the central level. Olfactory epithelia of the adult are located primarily on the third antennal segment (funiculus) and on the maxillary palps. About 200 basiconic (BS), 150 trichoid (TS) and 60 coeloconic sensilla (CS) cover the surface of the funiculus, and an additional 60 BS are located on the maxillary palps. Males possess about 30% more TS but 20% fewer BS than females. All these sensilla are multineuronal; they may be purely olfactory or multimodal with an olfactory component. Antennal and maxillary afferents converge onto approximately 35 glomeruli within the antennal lobe. These projections obey precise rules: individual fibers are glomerulus-specific, and different types of sensilla are associated with particular subsets of glomeruli. Possible functions of antennal glomeruli are discussed. In contrast to olfactory sensilla, gustatory sensilla of the imago are located at many sites, including the labellum, the pharynx, the legs, the wing margin and the female genitalia. Each of these sensory sites has its own central target. Taste sensilla are usually composed of one mechano-and three chemosensory neurons. Individual chemosensory neurons within a sensillum respond to distinct subsets of molecules and project into different central target regions. The chemosensory system of the larva is much simpler and consists essentially of three major sensillar complexes on the cephalic lobe, the dorsal, terminal and ventral organs, and a series of pharyngeal sensilla.     

The organization of the chemosensory system in Drosophila melanogaster: a rewiew

This review surveys the organization of the olfactory and gustatory systems in the imago and in the larva of Drosophila melanogaster, both at the sensory and the central level. Olfactory epithelia of the adult are located primarily on the third antennal segment (funiculus) and on the maxillary palps. About 200 basiconic (BS), 150 trichoid (TS) and 60 coeloconic sensilla (CS) cover the surface of the funiculus, and an additional 60 BS are located on the maxillary palps. Males possess about 30% more TS but 20% fewer BS than females. All these sensilla are multineuronal; they may be purely olfactory or multimodal with an olfactory component. Antennal and maxillary afferents converge onto approximately 35 glomeruli within the antennal lobe. These projections obey precise rules: individual fibers are glomerulus-specific, and different types of sensilla are associated with particular subsets of glomeruli. Possible functions of antennal glomeruli are discussed. In contrast to olfactory sensilla, gustatory sensilla of the imago are located at many sites, including the labellum, the pharynx, the legs, the wing margin and the female genitalia. Each of these sensory sites has its own central target. Taste sensilla are usually composed of one mechano-and three chemosensory neurons. Individual chemosensory neurons within a sensillum respond to distinct subsets of molecules and project into different central target regions. The chemosensory system of the larva is much simpler and consists essentially of three major sensillar complexes on the cephalic lobe, the dorsal, terminal and ventral organs, and a series of pharyngeal sensilla.     

Fine structure and primary sensory projections of sensilla on the maxillary palp of Drosophila melanogaster Meigen (Diptera : Drosophilidae)

Three types of hairs were identified on the maxillary palp of Drosophila melanogaster Meigen (Diptera : Drosophilidae): (i) single-walled, multiporous sensilla basiconica, which constitute 75% of the innervated hairs; (ii) thick walled non-porous sensilla trichodea, which make up the remaining 25% of the innervated hairs; and (iii) numerous spinules, which are un-innervated. These sensilla basiconica uniformly contain 2 bipolar sense cells, whereas sensilla trichodea have a single dendrite with a tubular body at the base of each hair. A majority of the sensilla basiconica is located on the distal half of the dorsal surface, whereas sensilla trichodea are positioned on the tip and entire ventrolateral ridge of the palp. Approximately 125 axons of the sense cells join to form a single nerve. The structure of sensilla basiconica and sensilla trichodea suggests that they are olfactory and mechanosensory respectively. The contact chemoreceptors (gustatory sensilla) are conspicuously absent on the maxillary palp.Golgi silver impregnations and cobalt fills show that the primary sensory fibres from sensilla trichodea and sensilla basiconica on the maxillary palp project in the posterior suboesophageal ganglion (SOG) and the antennal lobe respectively. A single fibre projects separately either in the SOG or in the antennal lobe. In the antennal lobe, the input received from sensilla basiconica is usually bilateral and at least 5 glomeruli are innervated symmetrically on either side from both the palps.This study suggests that the sensory neurons are capable of making selective projections in the specific regions of the brain. Accordingly, the fibres from a sensillum project to the brain with respect to their functions and the individual glomeruli represent functional units of the brain, receiving inputs in a characteristic combination.     

Ultrastructure of a possible new type of crustacean cuticular strain receptor in Carcinus maenas (crustacea,decapoda)

A hitherto unknown sensillum type, the “intracuticular sensillum” was identified on the dactyls of the walking legs of the shore crab, Carcinus maenas. Each sensillum is innervated by two sensory cells with dendrites of “scolopidial” (type I) organization. The ciliary segment of the dendrite is 5–6 μm long and contains A-tubules with an electron-dense core and dynein arm-like protuberances; the terminal segment is characterized by densely packed microtubules. The outer dendritic segments pass through the endo- and exocuticle enclosed in a dendritic sheath and a cuticulax tube (canal), which is suspended inside a slit-shaped cavity by cuticular lamellae. The dendrites and the cavity terminate in a cupola-shaped invagination of the epicuticle. External cuticular structures are lacking. Three inner and four to six outer enveloping cells are associated with each intracuticular sensillum. The innermost enveloping cell contains a large scolopale that is connected to the ciliary rootlets inside the inner dendritic segments by desmosomes. Scolopale rods are present in enveloping cell 2. Since type I dendrites and a scolopale are regarded as modality-specific structures of mechanoreceptors, and since no supracuticular endorgan is present, the intracuticular sensilla likely are sensitive to cuticular strains. The intracuticular sensilla should be regarded as analogous to insect campaniform sensilla and arachnid slit sense organs.     

Immunolocalization of general odorant-binding protein in antennal sensilla of moth caterpillars

Antennae of Bombyx mori and Helicoverpa armigera larvae were immunolabelled with antisera raised against the pheromone-binding protein or the general odorant-binding protein 2 of Antheraea polyphemus to assign the expression of these proteins to individual sensilla and to compare the localization pattern with that in sensilla of adult moths. Specific labelling of antennal sensilla was only obtained with the antiserum against general odorant-binding protein 2. Among the few sensilla present on the antenna the three large sensilla basiconica, which are suspected to be olfactory in function, were labelled. These sensilla are compound sensilla consisting of several sensillum units which form a common sensory hair. The hair is single-walled and pierced by many pores. Labelling of sensillum compartments was the same as in sensilla of adults. Prominent labelling of the sensillum lymph is accompanied by labelling of secretory organelles in the two outermost auxiliary cells and of endocytotic pathways in all sensillum cells. The results suggest that general odorant-binding protein is expressed in single-walled multiporous sensilla of presumed olfactory function on the antenna of moth larvae. The overall identity of the localization pattern for general odorant-binding protein between larval and adult sensilla implies a similar role of these proteins in olfactory stimulus transduction.