Morphology of the prothoracic discs and associated sensilla of Acanthocnemus nigricans (Coleoptera, Acanthocnemidae)

The Australian ‘little ash beetle’ Acanthocnemus nigricans (Coleoptera, Cleroidea, Acanthocnemidae) is attracted by forest fires. A. nigricans has one pair of unique prothoracic sensory organs and it has been speculated that these organs may play a role in fire detection. Each organ consists of a cuticular disc, which is fixed over an air-filled cavity. On the outer surface of the disc, about 90 tiny cuticular sensilla are situated. The poreless outer peg of a sensillum is 3–5 μm long and is surrounded by a cuticular wall. One ciliary sensory cell innervates the peg. As a special feature, the outer dendritic segment is very short already terminating below the cuticle. A massive electron-dense cylindrical rod, which most probably represents the hypertrophied dendritic sheath, extends through the cuticular canal connecting the tip of the outer dendritic segment to the peg. The dendritic inner segment and the soma are fused indistinguishably. Thin, leaflike extensions of glial cells deeply extend into that conjoint and considerably enlarged compartment which also contains large numbers of mitochondria. In summary, the sensilla of the sensory disc of A. nigricans represent a new type of insect sensillum of hitherto unknown function. The possible role of the prothoracic sensory organ in fire detection is discussed.     

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.     

Sensilla on the maxillary palps of Helicoverpa armigera caterpillars: in search of the CO(2)-receptor

The ultrastructure of sensilla on the maxillary palps of helicoverpa armigera caterpillars has been investigated in order ot find candidates for CO(2)-receptors. The following sensilla are found on the palps: a) 8 chemosensory pegs at the tip; b), a large distal pore plate; c), a smaller proximal pore plate; d), a digitiform organ; e), a campaniform sensillum; and f), 3 scolopidia. Each chemosensory peg at the tip is innervated by 4-5 sensory neurons. Five of these pegs are most probably contact chemoreceptors, because each has a dendrite with a tubular body. The distal pore plate has a porous cuticle and is innervated by 3 sensory neurons, each of which sends a highly branched dendrite into a large cuticular cavity. The proximal pore plate is made up from two fused organs, has also a porous cuticle, and is innervated by two sensory neurons which send their dendrites into a narrow cuticular channel. The digitiform organ is innervated by one sensory cell which sends a highly lamellated dendrite into a narrow channel within a chip-shaped protrusion of the porous cuticle. For several reasons, the digitiform organ is the most probable candidate for the CO(2)-receptor. Another possible candidate is the distal pore plate.     

Lamellated outer dendritic segments of a chemoreceptor within wall-pore sensilla in the labial palp-pit organ of the butterfly,Pieris rapae L. (Insecta,Lepidoptera)

Summary The structure of the sensilla in the apical pit of the third segment of the labial palps in Pieris rapae was investigated in cryofixed and chemically fixed specimens. There is a field of about 80 club-shaped sensilla, 94% of which house a single sensory cell; 6% contain two sensory cells. All sensory cells are of the same type and are characterized by the structure of the dendritic outer segment. This consists of a proximal cylindrical and a distal lamellated section. The lamellae contain a lattice of longitudinally arranged microtubules. Filamentous strands connect the microtubules with the surface membrane of the lamellae. The surface area of the lamellated section amounts to about 40 m². Pores and pore tubules are present in the cuticular wall of the peg. Electrophysiological recordings show that the sensory cells are olfactory receptors, which react to a variety of complex plant odors and to the odor of conspecifics. It is shown that (a) the usual modality-specific characteristics of insect olfactory sensilla apply here also; (b) lamellation is not only a characteristic of thermoreceptors, but also of olfactory chemoreceptors; (c) there are pore tubules that are separated from the dendritic membranes by an extended dendritic sheath; and (d) in the labial palppit sensilla only the lamellated dendritic tip region may be involved in sensory transduction.Supported by the Deutsche Forschungsgemeinschaft (SFB 4/G1)     

The fine structure of thermo-/hygrosensitive sensilla in the silkmoth Bombyx mori: Receptor membrane substructure and sensory cell contacts

Summary The thermo-/hygroreceptive sensilla styloconica of the silkmoth Bombyx mori are located on the tips of the antennal branches. A small poreless cuticular peg is innervated by three sensory cells. The outer dendritic segments of two type-1 receptor cells, the presumed hygroreceptors, almost completely fill the peg lumen and are in close contact with each other. The outer dendritic segment of the third (type-2) receptor cell, the presumed thermoreceptor, forms lamellae below the peg base. The membranes of these lamellae are studded with knobs in orthogonal array, protruding into the extracellular space with the same orientation on facing lamellae. This Bossy Orthogonal Surface Substructure (BOSS) is assumed to play a role in thermoreceptor function. Contacts are observed between the outer dendritic segments, between the inner dendritic segments immediately below the ciliary segments, and between the sensory cell somata. These contacts, which are not found in the olfactory sensilla (s. trichodea and basiconica) of this species, indicate electrical interactions between the three sensory cells of the styloconic sensillum and possibly are involved in the antagonistic and/or bimodal response characteristics of thermo-/hygroreceptor cells.     

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.     

Embryonic development and molting of the antennal coeloconic no pore- and double-walled wall pore sensilla in Locusta migratoria (Insecta,Orthopteroidea)

Summary The embryonic development of antennal coeloconic sensilla was studied at four stages between 132 and 252 h after oviposition in Locusta migratoria. Initially the anlagen of the sensilla consist of 2–4 sensory cells and 3 enveloping cells. Two additional cells contribute later to the formation of socket and pit. The dendritic outer segments of the sensory cells elongate before the trichogen process grows out (ecdysis type I) with exception of one sensory cell in anlagen of poreless (np) sensilla. Other differences between np and double-walled wall pore (dw wp) sensilla are not visible until at least about 220 h after oviposition. Molting, which was studied in four stages, follows ecdysis type I in both sensillum types. The fourth enveloping cell maintains its tight connection to the socket of the sensillum even after apolysis. Its apical portion is torn off and shed together with the old cuticle. The electron-dense material between the dendritic sheath and the cuticular wall of the peg in np sensilla, which is regarded important for stimulus transmission, is not deposited during retraction of the trichogen cell. The concentric walls and spoke channels characteristic of dw wp sensilla result from deposition of cuticular material around wedge-shaped projections of the trichogen cell. The typical trilaminar 15 nm cuticulin layer is produced only on the ridges of these sensilla. The first cuticular lining of the spoke channels is only 7 nm thick and of a different structure. A flocculent material surrounds the outgrowing trichogen process. It is continuous with the filling of the spoke channels and can thus be considered as component of the stimulus-transmitting material in the functioning intermolt dw wp sensilla.     

Lamellated outer dendritic segments of a sensory cell within a poreless thermo- and hygroreceptive sensillum of the insect Carausius morosus

Summary A sensillum in a narrow pit with a broad cuticular collar, located in a sensillum field on the 12th segment of the antennae of Carausius morosus, was investigated electrophysiologically. After marking, it was also examined with the transmission and the scanning electron microscopes. The number of sensory cells within the sensillum varies between three and four. One cell, present in half of the sensilla studied, exhibits a simple cilium of the 9×2+0 type as outer dendritic segment. The outer segment of a second unit is noteworthy in that it divides near its ciliary base into two branches. These flatten to form lamellae, then fold and wrap around each other. The remaining two sensory cells bear unbranched or bifurcate outer segments which contain densely packed microtubules. Only these outer segments extend into the cuticular peg; the others end beneath its base. The cuticular peg is devoid of pore systems. Electrophysiological recording yielded evidence that a cold, a dry and a moist air receptor are present. The fourth unit did not respond clearly to stimulation.Supported by the Deutsche Forschungsgemeinschaft (Al 56/6)Research Fellow of the Alexander von Humboldt Foundation     

Functional morphology of a double-walled multiporous olfactory sensillum: the sensillum coeloconicum of Bombyx mori (Insecta, Lepidoptera)

The fine structure of coeloconic sensilla of Bombyx mori was studied in cryofixed specimens. These sensilla belong to the category of double-walled wall-pore sensilla. The pegs are approximately 10 mum long, located in pits on the dorsal side of the antennal branches, and longitudinally grooved in their distal half (grooved surface approximately 30 mum(2)). The central lumen contains the outer dendritic segments of usually five receptor cells, and is surrounded by up to 15 partially fused cuticular fingers. The peripheral lumina of these cuticular fingers are filled with material resembling wax-canal filaments. Radial spoke channels ( approximately 600 per peg), each 10-20 nm wide, connect the central lumen with the longitudinal groove channels. Groove and spoke channels are assumed to mediate the transport of odorant molecules from the outer epicuticular surface layers to the sensory dendrites. Thus the double-walled wall-pore sensilla represent a bauplan essentially different from single-walled wall-pore sensilla; the reason, however, why the two types are found together throughout the insect orders remains enigmatic. Other peculiar features of the coeloconic sensilla of the silkmoth are invaginations of the outer dendritic segments and direct contacts between the receptor cell somata. The latter may be the structural correlate to electrophysiological observations indicative of peripheral interaction between the receptor neurons. All three auxiliary cells have elaborately folded apical plasma membranes studded with portasomes and associated with an abundance of mitochondria; basally they often contact tracheal branches. As compared to the auxiliary cells of the single-walled olfactory sensilla of the same species, all the mentioned features are much more prominent and hint to a higher ion pumping activity at the border to the sensillum-lymph cavities.     

The structure of bimodal chemo-, thermo-, and hygroreceptive sensilla on the antenna of Locusta migratoria

Summary Coeloconic sensilla on the antenna of Locusta migratoria were investigated electrophysiologically, labeled and then examined under the scanning or transmission electron microscope. They can be categorized into two main morphological types: 1) sensilla with wall pores and two concentric cuticular walls (wp-dw sensilla), and 2) sensilla lacking wall pores (np-sensilla).Hygroreceptors were observed only in np-sensilla. Olfactory receptors were observed in wp-dw sensilla, but not in np-sensilla. Cold-sensitive units were found in both types. Wp-dw sensilla with a cold-sensitive unit also manifest a fatty acid olfactory receptor. This cold-sensitive unit appears to be less sensitive to drops in temperature than the cold receptor combined with hygroreceptors in np-sensilla. Recordings from both types revealed up to three or sometimes four units that could be distinguished on the basis of impulse amplitude.The number of np-sensilla on each antennal segment ranges from 0 to 6. These sensilla are pegs 4–5 m long with a terminal swelling. The dendritic outer segments of two sensory cells fit tightly into these pegs. The dendrite of the third sensory cell ends at its base. All three are up to 1 m thick, unbranched, and densely filled with microtubules. The cuticular wall and socket of the peg are characterized by (1) an electron-dense inner layer surrounding the dendrites and continuous with electron dense clusters at the base of the peg, and (2) clefts in the cuticle at the insertion of the peg.The present observations favor the hypothesis that insect sensilla combining the triad of moist air, dry air and cold receptors generally lack pores. Specifical, though still alternative roles in stimulus transduction can be suggested for substructures demonstrated in the present and previous electron microscopic investigations.Supported by the Deutsche Forschungsgemeinschaft (SFB 4:G1/D)     

Ultrastructure and receptor cell responses of the antennal grooved peg sensilla of Triatoma infestans (Hemiptera: Reduviidae)

Ultrastructural examination of grooved-peg (GP) sensilla on the antenna of fifth instar Triatoma infestans nymphs by scanning electron microscopy and transmission electron microscopy reveal that they are 8–18 μm long with a diameter of about 2–2.8 μm at the non-articulated base. Some pegs have a terminal pore. These double-walled wall-pore (dw-wp) sensilla have an outer cuticular wall with 13–18 longitudinal grooves at the distal part of the peg. Groove channels are present at the bottom of the grooves from which radial spoke channels lead into the inner sensillum-lymph cavity. A dendrite sheath connects the tip of the thecogen cell to the inner cuticular wall thus forming separated outer and inner sensillum-lymph cavities. Four or five bipolar receptor cells are ensheathed successively within the GP sensilla by the thecogen cell, trichogen and tormogen cells. The inner dendritic segments of each sensory cell give rise at the ciliary constriction to an unbranched outer dendritic segment which can reach the tip of the sensillum.Electrophysiological recordings from the GP sensilla indicate that they house NH₃, short-chain carboxylic acid and short-chain aliphatic amine receptor cells and can be divided into three functional sub-types (GP 1–3). All GP sensilla carry a receptor cell excited by aliphatic amines, such as isobutylamine, a compound associated with vertebrate odour. GP type 1 and 2 sensilla house, in addition, an NH₃-excited cell whereas the type 2 sensilla also contains a short-chain carboxylic acid receptor. No cell particularly sensitive to either NH₃ or carboxylic acids was found in the grooved-peg type 3 sensilla. GP types 1, 2 and 3 represent ca. 36, 10 and 43% of the GP sensilla, respectively, whereas the remaining 11% contain receptor cells that manifest normal spontaneous activity but do not respond to any of the afore mentioned stimuli.     

Ultrastructural Analysis of the Sensilla of Austroperipatus aequabilis Reid, 1996 (Onychophora,Peripatopsidae)

The head of Austroperipatus aequabilis bears five types of sensilla. which were examined by electron microscopy. They differ from each other in position, shape of outer sensory elements and cuticular socket structures. Thus, we distinguish sensilla with sensory hairs, sensilla with sensory bulbs, cone-shaped sensilla. sensilla with sensory bristles, and sensilla of the lips. They are composed of up to 15 cells, which can he separated into four cell types. The most frequent cell type is the bipolar receptor cell that occurs in all sensilla. The apical surface of this primary receptor cell is characterized by one or two partly branched cilia with a basal 9 × 2 + 0 pattern of microtubules. A modified bipolar receptor cell was found in all sensilla bearing a sensory peg except for the sensilla equipped with sensory bristles. The apical dendrite extends to a long pale process which exclusively contains mitochondria and single microtubules. In all sensilla examined in this study at least one supporting cell occurs which is characterized by parallel microvilli. An additional function of this cell type as a part of the stimulus-conducting system is possible. In the sensillum with a sensory bulb two kinds of supporting cells occur. A unique cell type with an upside down position has regularly been found in all sensilla bearing a sensory peg. Apart from the sensilla they also occur within the labial epidermis. Since most sensilla contain several different receptor cells, they can be considered as complex sense organs. © 1998 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd. All rights reserved     

Development of antennal sensilla during moulting inNeomysis integer (Leach) (Crustacea,Mysidacea)

Summary The sensilla are associated with 6 enveloping cells. The innermost enveloping cell (e 1) secretes the dendritic sheath (=thecogen cell). All other enveloping cells are involved in the formation of the outer cuticular apparatus in secreting the cuticle of a definite region of the new hair shaft.The development of the new sensilla begins when an exuvial space expands between old cuticle and epithelium. The newly forming hair shafts lie folded back in an invagination of the epidermal tissue. Only a distal shaft part projects into the free exuvial space. The cuticle of the distal and middle shaft region is secreted by the three middle enveloping cells (e 2–e 4) (=trichogen cells), which are arranged around the dendritic sheath.The wall of the cylinder, in which the distal shaft is situated, is formed by the cuticle of the future proximal shaft region. It is secreted by the outer enveloping cells (e 5 and e 6). Furthermore, both enveloping cells form the hair socket (=trichogen-tormogen cells).The outer dendritic segments encased within a dendritic sheath run up through the newly formed hair shaft and continue to the old cuticular apparatus. The connection between sensory cells and old hair shaft is maintained until ecdysis. On ecdysis the old cuticle is shed and the newly formed shaft of the sensillum is everted like the invaginated finger of a glove. The dendritic sheath and the outer dendritic segments break off at the tip of the new hair shaft. Morphologically this moulting process ensures that the sensitivity of the receptors is maintained until ecdysis.The internal organization of the sensory cells shows no striking changes during the moulting cycle. An increased number of vesicles is accumulated distally within the inner dendritic segments and distributed throughout the outer segments of the dendrites. The cytoplasmic feature of the enveloping cells indicates that synthesis and release of substances for the cuticular apparatus of the new sensillum take place.     

Ultrastructure of the peg and hair sensilla on the antenna of larval Aedes aegypti (L.)

The antenna of fourth instar larvae of Aedes aegypti has one peg organ of a basiconic type innervated by four neurons. The dendrites are ensheathed to near their terminations at the peg tip by an electron-dense dendritic sheath and by a cuticular sheath. They have easy communication by diffusion with the external environment only at the tip through a peripheral ensheathing membrane and six slit-channels. One of the dendrites resembles a tubular body proximally and may be mechanoreceptive. The peg generally appears to be a contact chemoreceptor. There are three antennal hairs of a typical sensillum trichodeum type innervated at the base by one neuron each. An intricate terminal mechanism at the insertion of the dendrite in the hair is described. These are believed to be tactile hairs. There are also three antennal hairs each innervated by two neurons. The dendrite from one terminates at the base similar to that of a tactile hair, and is believed to function in a similar mechanoreceptive manner. The dendrite from the second neuron extends naked along the length of the hair lumen. It is believed to be primarily chemoreceptive, in a slow-acting general sensory function. In all the sensilla there appear to be secretions produced in the junction body regions of the dendrites, and there is evidence for accumulation of secretory materials in the dendritic tips in some of the sensilla.     

The thermo-/hygrosensitive sensilla of the silkmoth,Bombyx mori: morphological changes after dry- and moist-adaptation

Summary The thermo-/hygrosensitive sensilla styloconica of the silk moth Bombyx mori were studied using cryofixation and freez-substitution. These sensilla are characterized by a short poreless cuticular peg, which is double-walled in its distal part. The central lumen is innervated by the unbranched outer dendritic segments of the two presumed hygroreceptor cells. The presumed thermoreceptor cell displays lamellae below the peg base. Within the peg lumen, the dendrites are surrounded by the peridendritic dense coat and the lowdensity matrix. Below the peg base, these structures continue as the dendrite sheath, which is separated from the outer sensillum-lymph space by a layer of the trichogen cell. The central lumen, therefore, is only connected with the inner sensillum-lymph space, but the appearance of the low-density matrix, within the peg, differs from that of the sensillum lymph below the peg. In moist-adapted (24 h) sensilla, the two hygroreceptor dendrites invade the peg for three quarters and one half of its length, respectively, and fill the cross-sectional area of the lumen by 50–80%. In dry-adapted (24 h) sensilla, the dendrites terminate more proximally and fill the cross-section by 35%. The volume of the low-density matrix increases under dry conditions and decreases under humid conditions. At intermediate ambient humidity, the morphology of these sensilla is halfway between the dry-adapted and the moist-adapted state. The effect of dry-adaptation is reversible, so that sensilla that were first dry-adapted and then moist-adapted (24 h each) before cryofixation cannot be distinguished from moist-adapted sensilla. The reduction of the exposed length of the dendrites is interpreted as a shift of the working range of the receptors and/or protection against desiccation. The current theories of sensory transduction in hygroreceptors, in particular the hygrometer and evaporimeter hypotheses, are discussed with respect to the present findings.     

The cercal receptor system of the praying mantid,Archimantis brunneriana Sauss.

Summary The cerci of the praying mantid, Archimantis brunneriana Sauss., are paired segmented sensory organs located at the tip of the abdomen. Basally the cercal segments are slightly flattened dorso-ventrally and are fused to such a degree that it is difficult to distinguish them. Distally the segments become progressively more flattened laterally and their boundaries become more obvious.Two types of sensilla are present on the cerci, trichoid sensilla and filiform sensilla. Trichoid hairs are longest on the medial side of the cerci and toward the cercal base. On the proximal cercal segments they are grouped toward the middle of each segment while they are more uniformly distributed on the distal segments. Filiform sensilla are found at the distal end of each segment except the last and are most abundant on the middle segments of the cercus. Both the number of cercal segments and the number of sensilla are variable. Trichoid hairs are highly variable in appearance from short and stout to long and thin. They arise from a raised base, have a fluted shaft, and some have a pore at the tip. They are innervated by from one to five dendrites, one of which is always considerably larger than the others. Some of the dendrites continue out into the shaft of the hair.Filiform hairs have fluted shafts and are mounted in a flexible membrane within a cuticular ring in a depression. They are innervated by a single large sensory neuron, the dendrite of which passes across a flattened area on the inner wall of the lumen of the hair. The dendritic sheath forms the lining of the ecdysial canal and is therefore firmly attached to the hair. The dendrite is attached to the sheath by desmosomes distally and is penetrated by projections of the sheath more proximally. A fibrous cap surrounds the dendrite and may hold it in place relative to the hair.The cercal receptor system of Archimantis is compared to those of cockroaches and crickets.     

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.     

Labial tip sensilla of Blissus leucopterus leucopterus (Hemiptera: Blissidae): Ultrastructure and behavior

The cuticular sensory receptors that are found on the apex of the labium of hemipterans play an important role in their feeding behavior. In this study we describe the ultrastructure, number, and distribution of sensilla on the labium apex of the chinch bug, Blissus leucopterus leucopterus. Each apical field of sensilla on the labium contains 11 uniporous peg sensilla and one sensillum chaeticum. The uniporous peg sensilla are innervated by 4–5 bipolar neurons that send dendrites in the lumen of each peg. Three neurons are associated with each sensillum chaeticum, two neurons have dendrites in the lumen of the sensillum, and the third dendrite ends in a tubular body at the base of the sensillum. Behavioral tests that involve chemical blockage of the sensory receptors show the importance of the labial sensilla in feeding behavior. Both morphological and behavioral evidence indicate that the labial sensilla have a chemosensitive function.     

Fine structure of the pheromone-sensitive sensilla on the antenna of the hawkmoth, Manduca sexta

The flagellar antenna of the male hawkmoth Manduca sexta carries about 42,000 pheromone-sensitive sensilla trichodea, which are arranged in 'baskets' on the single segments. Each sensillum consists of a cuticular hair up to 500 mum long and is innervated by two bipolar sensory neurons. Each neuron sends an unbranched dendrite into the hair shaft. The dendrite is subdivided by a short ciliary region into an inner and an outer segment. The inner segment is especially rich in smooth vesicles, which accumulate beneath the ciliary region where they seem to fuse with the dendritic membrane. The outer dendritic segment often shows conspicuous 'beads' along its length. Three auxiliary, or enveloping, cells belong to each adult sensillum. These are the thecogen, the trichogen, and the 'outer' cell. Most probably, the latter is not homologous with the 'traditional' tormogen cell from a genealogical point of view.     

Fine structure and sensory apparatus of the mouthparts of the pear psyllid,Cacopsylla chinensis (Yang et Li) (Hemiptera: Psyllidae)

The pear psyllid, Cacopsylla chinensis (Yang et Li) (Hemiptera: Psyllidae), is one of the most significant economic pests of pear in China, causing direct damage through feeding by the highly specialized piercing–sucking mouthparts. The ultrastructural morphology and sensory apparatus of the mouthparts of the adult were examined using scanning and transmission electron microscopy. The piercing–sucking mouthparts of C. chinensis are composed of a three-segmented labium with a deep groove in the anterior side, a stylet fascicle consisting of two mandibular and two maxillary stylets, and a pyramid-shaped labrum. Proximal to the labium, the stylet fascicle forms a large loop within a membranous crumena. Mandibles, with more than ten teeth on the external convex region, can be seen on the distal extremity. Smooth maxillary stylets are interlocked to form a larger food canal and a smaller salivary canal. One dendritic canal housing 2 dendrites is also found in each mandible. Two types of sensilla trichodea, four types of sensilla basiconica, single as well as groups of sensilla campaniformia, and oval flattened sensilla occur in different locations on the labium, whereas a kind of sensilla basiconica is at the junction of the labrum and anteclypeus. Sensilla trichodea and sensilla campaniformia, always present with denticles, are present on the middle labial segment. Three types of sensilla basiconica, two types of sensilla trichodea and two oval flattened sensilla are located on the distal labial segment. The mouthpart morphology and abundance of sensilla located on the labium in C. chinensis are illustrated, along with a brief discussion of their taxonomic and putative functional significance.