Fine structure of antennal mechanosensilla of adult Rhodnius prolixus stål (Hemiptera: Reduviidae)

Each antenna of both sexes of adult Rhodnius prolixus has approximately 570 mechanosensitive neurons that innervate five morphologic types of cuticular mechanosensilla: campaniform sensilla, tapered hairs, trichobothria, and type I and type II bristle sensilla. Each campaniform sensillum and tapered hair is presumably innervated by one mechanosensitive bipolar neuron and probably functions in proprioception. The campaniform sensilla being located at the base of the scape could monitor the position of the antenna. Tapered hairs are found at the distal margin of flagellar segment I and projecting laterally from the bases of the pedicel and scape. They probably provide information about the relative positions of the antennal segments. Seven trichobothrium are located on the pedicel and three on flagellar segment I. Each trichobothrium has a long filamentous hair inserted into the base of a socket that extends inwardly as a cuticular tube and is innervated by one bipolar neuron with a tublar body, a parallel arrangement of microtubules associated with electron-dense material. The trichobothria may respond to small variations in air currents. Type I bristles occur at the base of the antenna and are the most numerous type of mechanosensillum; an average of 452 occur on each antenna of females and 440 on males. The bristle is curved toward the antennal shaft and is serrated distally. Type II bristles are located distally and are the second most numerous type of mechanosensillum; an average of 88 were counted on each antenna of females and 94 on males. The type II bristle is straight with small, longitudinal, external grooves and projects laterally from the antennal shaft. Each type I and II bristle sensillum is innervated by a bipolar neuron whose dendrite is divided into an inner and outer segment. The outer segment is encased by a dendritic sheath which may be highly convoluted and distally contains a tubular body. Two sheath cells are associated with each sensillum. Both types of bristle sensilla have a tactile function. The tubular bodies of both types of bristle sensilla have a complex structure indicating that they are very sensitive. Variations in the amount and arrangement of the electron-dense material at the tip of the tubular bodies may reflect differences in viscoelastic properties that underlie functional characteristics.     

Sensory cell degeneration in the ontogeny of the chemosensitive sensilla in the labial palp-pit organ of the butterfly,Pieris rapae L. (Insecta,Lepidoptera)

Summary The ontogeny of the chemoreceptive sensilla in the labial palp-pit organ was studied in Pieris rapae by examining twelve successive stages between pupation and emergence of the imago, which takes a period of 160 h under the experimental conditions. Mitoses occur until 20 h after pupation. They lead to anlagen of sensilla, 91% of which are comprised of three sensory cells. However, two sensory cells degenerate in each sensillum during a period of 28 h. The same process occurs in anlagen with four sensory cells resulting in bicellular sensilla. Axons grow out only after the number of sensory cells has been reduced. Further consecutive steps in sensory cell differentiation are: (a) outgrowth of dendritic outer segment and dendrite sheath; (b) outgrowth of trichogen process and change in structure of elongating dendrite sheath; (c) deposition of cuticle and pore tubules in the pegs; (d) retraction of trichogen process; (e) increase in diameter of dendritic outer segment accompanied by increase of microtubule number and appearance of regularly spaced electron-dense bodies at tubular doublets; (f) branching of dendritic outer segment; and (g) transformation of the dendritic branches into curled lamellae and partial destruction of the dendrite sheath. The unique process of sensory cell degeneration is interpreted as an event that revokes a step towards a possible functional improvement of the labial palp-pit organ during further evolutionSupported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

The morphology of spider sensilla. I. Mechanoreceptors

The common tactile hair sensilla of spider tarsi were studied in web spiders (Araneus) and ground spiders (Lycosa, Dugesiella) using scanning and transmission electron microscopy. All of these sensilla are innervated by three bipolar neurons whose dendrites end proximally at the sensillum base. Each dendritic terminal exhibits a tubular body, a dense array of microtubules typical for mechanoreceptive sensilla. A dendritic sheath encloses the outer dendritic segments and connects the dendritic terminals to cuticular components of the hair sensillum in three different ways: (1) A distal extension of the dendritic sheath connects to the midline of the hair base; (2) A forked arrangement of cuticular (?) strands attaches on both lateral sides of the hair base, and (3) The socket cuticle directly contacts a part of the dendritic sheath. The latter connection provides a fixed position for the three dendritic terminals and any movement of the hair shaft could be transmitted via connections (I) and (2). The triple innervation strongly suggests a directional sensitivity of these sensilla.Structural comparison between arachnid and insect mechanoreceptive sensilla indicates that tactile hair sensilla in Arachnida are multi-innervated whereas the corresponding reccptors in Insecta are singly innervated.     

Die digitiformen Sensillen auf dem Maxillarpalpus von Coleoptera

Summary The digitiform sensilla on the distal segment of the maxillar palps ofAgabus bipustulatus (L.) andHydrobius fuscipes (L.) were investigated by electron microscopic methods. Each sensillum is innervated by a single bipolar sensory cell. The sensilla ofHydrobius are associated with three enveloping cells, which enclose an inner and outer receptor lymph cavity. A single enveloping cell only is found in the completely differentiated sensilla ofAgabus. These sensilla do not form an outer lymph cavity. The area beneath the hair base is filled by the distal process of the enveloping cell and by extensions of epidermal cells. Only one extra-cellular space exists, which seems to be homologous to an inner receptor lymph cavity.The outer dendritic segment surrounded by a dendritic sheath runs to the tip of the hair shaft. In the hair shaft the outer dendritic segment divides into several branches. The poreless hair shaft does not rise over the surface of the cuticle, but it is positioned in a narrow shallow groove. Special socket structures or a tubular body do not exist. The digiti-form sensilla possess neither the typical feature of mechanosensitive, nor gustatory or olfactory sensilla. The functional significance of the structural divergences in the sensilla of both species and the presumed function of the sensilla are discussed referring to hygro- and thermo-receptors.

Unserem verehrten Lehrer, Herrn Prof. Dr. H.Risler, dem wir für vielfache Förderung danken möchten, zum 65. Geburtstag gewidmet.     

Fine structure of maxillary sensilla of larval Toxorhynchites brevipalpis (Diptera: Culicidae) with comments on the role of sensilla in behavior

Each maxilla of fourth instar Toxorhynchites brevipalpis bears nine sensilla: Four are located at the tip of the maxillary palp and five on the maxillary body. At the palp tip are three tapered pegs on bulbous bases (MS₁, MS₂, MS₆) that are innervated by four, two, and two neurons, respectively, and probably function in chemoreception. Also at the palp tip is a sturdy, cuticular rod with a lumen (MS₅) that opens distally to the exterior. The proximal end of the rod is closed by a cuticular base to which a single unbranched dendrite containing only a few microtubules is attached. The function of MS₅ is enigmatic; possibilities include mechanoreception and detection of infrared radiation. On the maxillary body are two tapered pegs on a common bulbous base (GS₁, GS₂) that are each innervated by three neurons, and probably are chemosensory. Three setae also occur on the maxillary body. They arise from prominent sockets and are each innervated by a neuron terminating at the hair base as a tubular body, a characteristic of cuticular mechanosensilla. The maxillary sensilla are innervated by a total of 18 neurons: 14 are probably chemosensory, three mechanosensory, and one is of unknown function. These results, combined with those from a previous study on antennal sensilla (Jez and McIver, '80), indicate that the mechanosensitive neurons of the antennae and maxillae are a relatively small percentage of the total mechanosensilla on the entire larva. In contrast the chemosensitive neurons of the antennae and maxillae provide most of the information about the chemical environment of the larva. T. brevipalpis has three less than the maximum of seven maxillary palpal sensilla found in larval mosquitoes so far studied. This difference may reflect a lesser need for sensory information about the acceptability of potential food in predators compared to browsers and filter-feeders.     

Fine structure of the galeal styloconic sensilla of larval Lymantria dispar (Lepidoptera: Lymantriidae)

Lepidopteran larvae possess two pairs of styloconic sensilla located on the maxillary galea. These sensilla, namely the lateral and medial styloconic sensilla, are each comprised of a smaller cone, which is inserted into a style. They are thought to play an important role in host-plant selection and are the main organs involved in feeding. Ultrastructural examination of these sensilla of fifth instar Lymantria dispar (L.) larvae reveal that they are each approximately 70 um in length and 30 um in width. Each sensillum consists of a single sensory peg inserted into the socket of a large style. Each peg bears a slightly subapical terminal pore averaging 317 nm in lateral and 179 nm in medial sensilla. Each sensillum houses five bipolar neurons. The proximal dendritic segment of each neuron gives rise to an unbranched distal dendritic segment. Four of these dendrites terminate near the tip of the sensillum below the pore and bear ultrastructural features consistent with contact chemosensilla. The fifth distal dendrite terminates near the base of the peg and bears ultrastructural features consistent with mechanosensilla. Thus, these sensilla each bear a bimodal chemo-mechanosensory function. The distal dendrites lie within the dendritic channel and are enclosed by a dendritic sheath. The intermediate and outer sheath cells enclose a large sensillar sinus, whereas the smaller ciliary sinus is enclosed by the inner cell. The neurons are ensheathed successively by the inner, intermediate, and outer sheath cells.     

Are the most numerous sensilla of terrestrial isopods hygroreceptors? Ultrastructure of the dorsal tricorn sensilla of Porcellio scaber

The ultrastructure of the tricorn sensilla of the woodlouse Porcellio scaber was investigated in cryofixed and freeze-substituted, or chemically fixed specimens. The tricorn sensilla have a foramenized triangular-shaped outer hair and bear a poreless rod-like inner hair. The conical base of the inner hair is connected to the base of the outer hair by a complex cuticular structure. Each sensillum contains three sensory cells. The tip of one of the three dendrites contains a tubular body and is clamped between two bulges of the dendritic sheath. The two other dendrites protrude to the tip of the inner hair, flush against the cuticular wall. The microtubules in the ciliary segments are arranged in nine double tubuli that have neither osmiophilic cores nor arms. The ciliary rootlets are small. The inner segment of the largest dendrite wraps around the two smaller dendrites and one of seven enveloping cells in a mesaxon-like manner. Although this ultrastructure deviates considerably from most crustacean mechanosensitive sensilla, it nevertheless suggests a mechanosensitive function, at least for one of the sensory cells. In many aspects, the tricorn sensilla resemble the thermohygrosensilla of insects. However, our results suggest that the structural criteria for thermo-hygro-sensitivity used in insects cannot simply be applied to crustaceans.     

Prey finding behavior and mechanosensilla of larval Toxorhynchites brevipalpis Theobald (Diptera : Culicidae)

Behavioral experiments demonstrated that starved 3rd-instar Toxorhynchites brevipalpis (Diptera : Culicidae) will attack a glass probe in response to vibrations alone. Frequencies in the range of 80–200 Hz elicited 85% of the attacks. The observation that most attacks occurred while the larva was drifting forward towards the probe, substantiates that the predatory behavior of these organisms is basically opportunistic.The main setae on the thorax and abdomen arise from setal support plates and are of 4 general morphological types: dendritic, smooth, spinulate spiniform, and unbranched aciculate setae. The numbers and lengths of the setae on 3rd- and 4th-instar larvae are given. Each seta is innervated by a bipolar neuron with a tubular body in the dendrite, a characteristic of mechanosensilla. By virtue of their abundance and location, it seems probable that one or more types of these setae sense the water vibrations capable of eliciting an attack response.The tubular bodies of the 4 types of setae are somewhat unusual in that they lack electron-dense material, except near sites of attachment of the microtubules to the dendritic membrane, yet possess a large number of complexly arranged microtubules. Among the setal types, variations were observed in the prominence of microtubular attachment sites, proportion of tubules associated with the sites, and orientation of microtubules. In spinulate spiniform setae, the arrangement of microtubules varies within the same tubular body; peripherally most of the microtubules are in one direction with little convergence and have only one distinct attachment site, whereas medially they converge considerably resulting in 2 groups, each associated with its own well-defined site.The attachment sites, which presumably through linkage with the dendritic membrane provide a means of initiating depolarization, are associated with the distal ends of almost all of the microtubules. This suggests that in these tubular bodies depolarization may be initiated through mechanical force acting along the length of the microtubules, that is, stretching and/or compression.     

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.     

Fine structure and role in behavior of sensilla on the terminalia of Aedes aegypti (L.) (Diptera: Culicidae)

The terminalia of male and female Aedes aegypti (L.) bear numerous hairs of various shapes and lengths, all of which are mechanoreceptors. Each hair is innervated by one bipolar neuron which contains ciliary rootlets, two basal bodies, and a region assuming the structure of a non-motile cilium. At the distal tip of the dendrite is a tubular body, a characteristic of cuticular mechanoreceptors. Covering the outer dendritic segment is a cuticular sheath which ends proximally in a net-like felt-work and distally attaches to the hair base. Each hair sensillum has two sheath cells. Presumed efferent fibers are associated with the sheath cells. On the insula of the female terminalia are a few campaniform sensilla, the domes of which are raised into small pegs. The sensilla on the terminalia function in copulation and oviposition and probably in warning. A sequence of neurological events is traced for copulation and oviposition. Other cuticular structures, viz., scales, microtrichia, acanthae, and aedeagal spines, which occur on the terminalia are not innervated.     

Sensitivity of an insect mechanoreceptor during moulting

ABSTRACT. The fine structure and the behavioural threshold for vibration sensitivity of the eight thoracic filiform hairs of Barathra brassicae caterpillars were investigated through an intermoult/moult cycle. Associated with each filiform hair is one bipolar sensory cell and three enveloping cells. The outer dendritic segment terminates in an ecdysial canal in the hair base and a tubular body lies at its distal end. Shortly before apolysis the dendrite elongates. By this means the connection between the sensory cell and the old cuticular apparatus is maintained while the epithelium and the old thoracic cuticle are separating. The new cuticular apparatus of the filiform hair is formed in the second half of the larval stage by the three enveloping cells. A second tubular body in the elongated outer dendritic segment is formed at the base of the replacement hair 10 h before next ecdysis, so that the new hair functions as soon as ecdysis is completed, the old cuticular apparatus with the old tubular bodies being torn away with the exuvia during ecdysis. Sensitivity to a 300 Hz tone was tested in the standing wave of a Kundt's tube. Throughout most of the larval instar the threshold was 2.0 ± 0.3 μm particle displacement amplitude until 1–2h before ecdysis when it rose to 6.8 ± 1.3 μm and at 10–30 min before the beginning of ecdysis no reaction to sound could be detected. Once the old cuticle was shed maximum sensitivity returned as soon as the replacement hairs were erect. The sensilla are therefore physiologically functional at all developmental stages except for 30–60 min during actual ecdysis.     

The mechanism of sensory transduction in the sensilla of the trochanteral hair plate of the cockroach,Periplaneta americana

Summary The trochanteral hair plate of the cockroach leg contains approximately 60 hair sensilla that are deflected by a joint membrane during flexion of the leg. Previous work has shown that the organ is a mechanoreceptor which limits leg flexion during walking by reflex connections to flexor and extensor motoneurons. Functional analysis of the largest sensilla has shown that their behaviour may be well approximated by a velocity detector followed by a unidirectional rectifier.We report here the results of an examination of the largest sensilla by scanning and transmission electron microscopy in an attempt to correlate the structure with the known functional elements. Each hair is innervated by a single sensory dendrite which is surrounded by an electron dense dendritic sheath. The dendrite terminates below the hair shaft in a tubular body containing a parallel array of microtubules embedded in an electron dense matrix, while the dendritic sheath extends beyond the tubular body to form the walls of the ecdysial canal. At the proximal end of the tubular body the dendritic sheath and sensory dendrite are anchored to the cuticular socket by a fibrous dome which seems to form a fulcrum around which the tubular body can be deflected by movements of the hair. We suggest that the basis for the detection of velocity may be mechanical differentiation by a fluid space between the dendritic sheath and the tubular body. The structure is also discussed with relation to the mechanism of sensory transduction and the possible causes of the unidirectional sensitivity.Supported by the Canadian Medical Research Council. The authors gratefully acknowledge the expert technical assistance of Sita Prasad     

Ultrastructure of the dendritic outer segments of sensory cells in poreless (‘no-pore’) sensilla of insects

Summary Poreless sensilla with inflexible sockets in insects presumably house hygro- and thermoreceptors (type-1, type-2 receptors). The dendritic outer segments of these receptor cells were studied mainly in cryofixed antennae of two species of moth (Antheraea pernyi, A. polyphemus) and one beetle (Aleochara curtula). As a rule two type-1 receptor cells are present. Their dendritic outer segments do not branch. They project into the distal cuticular parts of the sensillum and are in close contact with its four-layered wall. The segments differ in shape and microtubule density. As well, in A. curtula, the microtubules are interconnected by electron-dense material for some distance, thus forming a tubular body-like structure of 1.3 m length. The dendritic outer segment of the single type-2 receptor cell is branched and lamellated. Its lamellae are connected by structures similar to septate junctions, which occupy about 70% of the total surface of the lamellated portion of the dendrite. In tangential sections, the septa appear as parallel strands approximately perpendicular to the long axis of the dendritic segment. The structure of type-1 receptors is discussed with regard to the hypothesis for a mechano-electrical transduction. The possible functions of lamellation and junctional connections in type-2 receptors are discussed.Supported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

Structure and differentiation of the sensilla of the ventral sensory field on the maxillary palps ofPeriplaneta americana (Insecta,Blattodea), paying special attention to the ciliogenesis of the sensory cells

Summary The embryonic development of palpal contact chemosensitive sensilla was studied from 42% of development up to the hatching of the larvae. Ciliogenesis of the sensory cells can be observed at the earliest stages investigated. A complex consisting of two basal bodies and a cap-like ciliary vesicle is localized in the dendritic inner segment. It migrates apically and fuses with the cytoplasmic membrane. At the same time, microtubule doublets of the distal basal body elongate, thus generating the dendritic outer segment. Furthermore, the typical accessory structures of a motile cilium are formed. Although the central pair of microtubules is lacking, the dendritic outer segment can be considered as a modified motile cilium. At about 84% of development the hair structure starts to be formed. Whereas the socket is generated by the tormogen cell, the trichogen cell produces the hair shaft and terminal porus. The dendrite sheath, which rises above the newly formed hair, is attached apically to the embryonic cuticle forming an irregular pore. In larvae and imagines, the inner surface of the dendrite sheath is highly differentiated. A range of circular ledges and filamentous structures wrapping around the dendritic outer segments can be distinguished. These may have a stabilizing function. Furthermore, in cryofixed specimens, the dendritic outer segments possess regularly spaced swellings which are about 1 m in length and about 0.5 m in diameter. Their functional significance is still unclear.     

Comparison of the ultrastructure of stimulated and unstimulated mechanoreceptors in the taste hairs of the blowfly Phaenicia serricata

The structure of mechanoreceptors at the base of labeilar taste hairs of the blowfly Phaenicia serricata were examined in stimulated and unstimulated conditions (i.e. with the hair bent or unbent). Physiological recordings from the mechanoreceptor showed that the receptors responded when the hair is bent dorsally or ventrally and when the hair is bent at extreme angles. These conditions are the same as those placed on hairs in the anatomical studies. Bending the hair toward the ventral labellar surface caused the hair base to compress and indent the tubular body and its surrounding membrane and sheath at the distal end of the mechanoreceptor dendrite. In compressed tubular bodies, microtubules oriented longitudinally were bent and separated a greater distance from each other. Separation as much as 70 nm was observed in compressed tubular bodies as compared with a maximum of 26 nm between microtubules in tubular bodies of unbent hairs. The dense amorphous material between microtubules of compressed tubular bodies formed prominent bridges 18 nm thick connecting the microtubules at intervals of 48–74 nm. Thin 10 nm filaments were also evident in the spaces between microtubules. When the hair was bent toward the proximal end of the proboscis, the tip of the tubular body was bent about 15 °. The tubular body appears to function as a firm but resilient structure over which the dendritic membrane can be stretched during mechanostimulation. Comparison of morphology of bent and unbent hairs suggests a means by which mechanical force from the movement of the hair is transferred to the receptors by structures in the hair socket region. No differences were found in ciliary structures of stimulated and unstimulated receptors.     

Morphology and ultrastructure of the sensory spine,a presumed mechanoreceptor of Sphaeroma hookeri (Crustacea,Isopoda), and remarks on similar spines in other peracarids

The isopod Sphaeroma hookeri and many other isopods and peracarids have a sensory spine with laterally inserting sensory hair, positioned in the apical region of the propodal palm of pereopod 1. This spine is innervated by five to eight sensory cells (each giving rise to one cilium) the dendrites of which can be divided into an inner and outer dendritic segment. The cilia are surrounded by an extracellular, electron-dense dendritic sheath. Thirteen enveloping cells are present. The outer dendritic segment (structure beyond the basal bodies) contains two receptor lymph cavities; the inner one lying within the dendritic sheath is homologous with the inner receptor lymph cavity of insects. Scolopales, or tubular bodies, are lacking; their function is probably accomplished by the dendritic sheath. Apically the sensory hair does not have a pore, and the spine is heavily sclerotized. The inner dendritic segment begins with a basal body from which rootlets of different length and thickness extend into the dendrite. In the latter is an accumulation of vesicles. The dendrites keep close contact with other dendrites and the enveloping cells by desmosomal membrane structures. The possible importance of the sensory spine for phylogenetic studies is discussed.     

Hair regeneration during moulting in the spiderCiniflo similis (Araneae,Dictynidae)

Summary The mechanoreceptive and chemoreceptive hairs on the legs of the cribellate spiderCiniflo similis were examined during the moulting cycle. In mechanoreceptive hairs the new hair shaft is formed around the extended dentrites, which emerge from near the tip of the newly forming hair and continue to the old sensillum within the extended dendritic sheath. Thus there is no ecdysial canal in the base of the hair shaft as found in insect mechanoreceptive hairs. The dendritic connection with the old hair is maintained until shortly before ecdysis by which time new tubular bodies have developed in the same dendrites at the base of the new hair. In chemoreceptive sensilla the new hair shaft is also formed around the elongated outer segment of the dendrites (19 chemosensitive and 2 mechanosensitive). The two mechanosensitive dendrites develop new tubular bodies at the base of the hair. As ecdysis occurs the old dendritic sheath and dendrites are snapped off at the tip of the new hair but the pore remains open. The ultrastructural evidence indicates that the roles of the three main enveloping cells are as follows: The dendritic sheath cell secretes the dendritic sheath, the middle enveloping cell forms the hair shaft while the outer enveloping cell forms the socket. This pattern corresponds closely to that observed in insecta sensilla. The extreme length of the chemoreceptive dendrites during moulting is mentioned in connection with receptor function. The unique multi-layered nature of the middle enveloping cell is seen as a device for the formation of regularly occurring rows of small spines on the shaft of the hair.     

Antennal sensilla ofNeomysis integer (leach)

Summary The most frequent type of the hair sensilla on the antennae ofNeomysis integer is investigated by electron microscopic methods. The cellular properties of the sensilla are compared with those of other arthropods in order to detect possible homologies.The hairs are innervated by 2, 3, 6, 8, 9, or 10 sensory cells. The dendrites show an inner and outer dendritic segment. Five or six enveloping cells belong to a sensillum. In intermoult stage, processes of all the enveloping cells except the innermost one extend into the hair shaft. The sensory hairs possess only a single liquor cavity, which morphologically is homologous to the inner lymph cavity of insect sensilla. Around the liquor cavity, a supporting structure is located which seems to be identical to the scolopale of chordotonal organs. The six-to tenfold-innervated hairs possess two groups of differently structured dendrites which are regularly arranged on opposite sides of the liquor cavity. The outer dendritic segments are enclosed in a dendritic sheath. It is secreted by the innermost enveloping cell (= dendritic sheath cell of insect sensilla). All the outer dendritic segments terminate in the distal region of the hair shaft which shows a pore at its tip. The possible function of the sensilla is discussed. The double and triple-innervated hairs are considered to be mechano-receptors, whereas the sensilla associated with six to ten sensory cells might be mechano-chemoreceptors.     

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.     

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.