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.     

A new mechanosensory organ on the anterior spinnerets of the spiderCupiennius salei (Araneae,Ctenidae)

We describe hitherto unknown mechanoreceptors on the anterior spinnerets of the spiderCupiennius salei. These receptors are found at the base of the spigots of the major ampullate glands which produce the dragline used by the spider as a safety thread in various behavioral situations. There are 40–60 mechanoreceptors associated with two spigots of each anterior spinneret. They are likely to provide information on the forces pulling on the dragline and also on its orientation in space. A single sensillum consists of a hole in the cuticle covered by a thin cuticular membrane. It much resembles spider slit sensilla, which are known to detect strains in the exoskeleton. Each sensillum is supplied by two dendrites most likely belonging to two bipolar sensory cells. One of the dendrites ends at the covering membrane and the other more proximally. The sensilla are arranged with their long axes roughly parallel to the circumference of the spigots. External forces, transmitted by the dragline, result in deformation of the central part of the cuticular plate at the base of the spigots and thus in stimulation of the sensilla. This is shown electrophysiologicallly. Considering their morphology, topography, and electrophysiology, these mechanoreceptors are suggested to be important in the sensory control of dragline release by the spider.     

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.     

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.     

The morphology and physiology of CAP organs in Jasus novaehollandiae (crustacea,decapoda, reptantia,macrura)

The location and external anatomy of the CAP organs of Jasus novaehollandiae were examined and found to be similar to those in Homarus gammarus (Laverack, 1978a). Histological examination of the organs showed threads or filaments arising from the internal surface of the spines to run through canals in the cuticle and join with dendrites of CAP sensory cells in the region of the hypodermis. The CAP neuron cell bodies lie in the nearby chordotonal organ strand. It is demonstrated that flexion of the limb causes the articulating membrane to deflect the spines of the sensillae distally. A variety of experimental techniques used to investigate the physiology of the organs reveals why previously reported attempts to record from the receptors failed. Direct stimulation of the sensillae with an analogue of the membrane and summation of many traces revealed phase constant responses at points corresponding to the covering and uncovering of the sensillae.     

Structure and localization of sensilla on antennae of caddisflies (Insecta: Trichoptera)

Structure of antenna segments and ultrastructure of antennal sensillae in representatives of 28 caddisfly families from both modern suborders were studied by electron and light microscopy methods. On Trichoptera antennae, 16 types of sensillae were found, some of them being described for the first time. Morphological peculiarities of cuticular ultrastructures on the antennal surface demonstrate essential differences in structure both at the family level and at the lower taxonomic levels. Specialized sensory fields structurally different from the remaining antennal surface were revealed on antennal flagellae in representatives of the Phryganeina suborder. A new classification of sensillae based on the structure of their cuticular section is proposed.     

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.     

The coelocapitular sensillum,an antennal hygro- and thermoreceptive sensillum of the honey bee,Apis mellifera L.

Summary The sensillum coelocapitulum, a hygro- and thermoreceptive sensillum of the honey bee, Apis mellifera, was investigated by electron microscopy. The cuticular apparatus of the sensillum is a mushroomshaped protrusion, devoid of pores, set in a narrow cylindrical pit positioned centrally within a cuticular, shallow depression. There may be three or four receptor cells. Three receptor cells have unbranched sensory cilia, containing densely packed microtubules, which extend distally into the cuticular apparatus and completely fill its cavity. These connecting cilia are of the usual 9+0 type. The fourth receptor, if present, has a thin sensory cilium which terminates beneath the cuticular apparatus. Its connecting cilium has armed outer doublets. The outer cavity is formed by two enveloping cells and is completely sealed off. Lipid deposits are present within the cavity and the tormogen cell. The thecogen cell has scolopale rod-like structures around the inner cavity. Features common to the insect hygro- and thermoreceptive sensilla are discussed in comparison with those of other insects.     

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

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

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.     

Spike amplitude of single-unit responses in antennal sensillae is controlled by the Drosophila circadian clock

Circadian changes in membrane potential and spontaneous firing frequency have been observed in microbial systems, invertebrates, and mammals. Oscillators in olfactory sensory neurons (OSNs) from Drosophila are both necessary and sufficient to sustain rhythms in electroanntenogram (EAG) responses, suggesting that odorant receptors (ORs) and/or OR-dependent processes are under clock control. We measured single-unit responses in different antennal sensillae from wild-type, clock mutant, odorant-receptor mutant, and G protein-coupled receptor kinase 2 (Gprk2) mutant flies to examine the cellular and molecular mechanisms that drive rhythms in olfaction. Spontaneous spike amplitude, but not spontaneous or odor-induced firing frequency, is under clock control in ab1 and ab3 basiconic sensillae and T2 trichoid sensillae. Mutants lacking odorant receptors in dendrites display constant low spike amplitudes, and the reduction or increase of levels of GPRK2 in OSNs results in constant low or constant high spontaneous spike amplitudes, respectively. We conclude that spike amplitude is controlled by circadian clocks in basiconic and trichoid sensillae and requires GPRK2 expression and the presence of functional ORs in dendrites. These results argue that rhythms in GPRK2 levels control OR localization and OR-dependent ion channel activity and/or composition to mediate rhythms in spontaneous spike amplitude.     

Fine structure of the chemoreceptor sensillum in Limulus

Each chemoreceptor sensillum of Limulus polyphemus consists of 6–15 bipolar neurosensory cells with distal processes confined within a single cuticular tubule as they extend to the outside environment. The cuticular tubule, which is enveloped by the cuticulo-tubal cell, opens proximally into a fluid-filled extracellular space through which the dendrite passes before entering the cuticular tubule. Between the neurosensory cells are one to three microvillar cells also exposed to the extracellular space. This space is enclosed by a sheath cell extending proximally from the inner opening of the cuticular tubule and enveloping the proximal portions of the dendrites, the distal portions of the microvillar cells, as well as the distal portion of some neurosensory cell bodies. Most of the remaining portions of the neurosensory cells and microvillar cells are enveloped by neuroglia. Tight junctions occur between the distal portions of the dendrites in or near the cuticular tubule. Each dendrite has a cilium-like segment located where it traverses the extracellular space with a 9 + 0 pattern of fibers. Septuplelayered junctions occur among the proximal portions of some dendrites and some neurosensory cell bodies of the same sensillum. The subjacent processes of the sensillum frequently course proximally as isolated axons before joining nerve bundles. In the chilarial and gnathobasal chemoreceptors these nerve bundles course proximally to neuropile clumps of a peripheral nerve plexus. The presence of numerous synaptic vesicles in the neuropiles suggests that chemical transmission may occur among “en passant” synapses formed by the axons. Proximally the neuropiles are joined to the central nervous system by relatively long nerves.     

Gill receptor arrays in the horseshoe crab (Limulus polyphemus)

The branchial warts on the endopodites of the gills are covered with goblet-shaped cuticular appendages, whose internal structure shows them to be chemoreceptors. The innervated goblets have a cuticular tubule that connects an external pore through their hollow interior with the epidermal sensillum. Associated sensory neurons give rise to small axons that pass through a synaptic plexus below the epidermis. The sense organs seem specialized for sampling the exhalent water current.     

Sensory activity affects sensory axon development in C. elegans

The simple nervous system of the nematode C. elegans consists of 302 neurons with highly reproducible morphologies, suggesting a hard-wired program of axon guidance. Surprisingly, we show here that sensory activity shapes sensory axon morphology in C. elegans. A class of mutants with deformed sensory cilia at their dendrite endings have extra axon branches, suggesting that sensory deprivation disrupts axon outgrowth. Mutations that alter calcium channels or membrane potential cause similar defects. Cell-specific perturbations of sensory activity can cause cell-autonomous changes in axon morphology. Although the sensory axons initially reach their targets in the embryo, the mutations that alter sensory activity cause extra axon growth late in development. Thus, perturbations of activity affect the maintenance of sensory axon morphology after an initial pattern of innervation is established. This system provides a genetically tractable model for identifying molecular mechanisms linking neuronal activity to nervous system structure.     

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.     

Ultrastructure des sensilles trochanteriens de Campodea sensillifera conde et mathieu (Diplura : Campodeidea)

The fine structure of the basiconica sensilla situated on the posterior part of trochanters in Campodea sensillifera (Diplura : Campodeidea) reveals that they are probably olfactory and mechano-sensitive setae. Each sensillum is composed of one sensory axis made of 3 dendrites ensheathed by 3 cells (thecogen, trichogen and tormogen); one outer segment ends by a tubular corper without connection with the cuticular layer. The setae are generally racket-shaped. The epicuticular layer of the expanded part is perforated by a lattice of numerous slits, which communicate with underlying canals. The ciliary structures and apex of the tormogen cell are eliminated just before ecdysis. The ciliary microtubules are present in the cavity of the new sensillum, but after ecdysis the microtubules persist only at the lower part of the peduncle. An ecdysial canal appears at the tip of the sensillum.     

Fine structure and distribution of antennal sensilla of the desert locust, Schistocerca gregaria (Orthoptera: Acrididae)

The fine structure and distribution of various types of antennal sensilla in three nymphal stages and in adults of both solitary-reared (solitary) and crowd-reared (gregarious) phases of the desert locust, Schistocerca gregaria, were investigated by scanning and transmission electron microscopy. Four types of sensilla were identified: sensilla basiconica, s. trichodea, s. coeloconica and s. chaetica. S. basiconica contain up to 50 sensory neurons, each of which displays massive dendritic branching. The sensillar wall is penetrated by a large number of pores. In contrast, s. trichodea contain one to three sensory neurons that branch to give five or six dendrites in the sensillar lumen; the sensillum wall is penetrated by relatively few pores. The s. coeloconica are situated in spherical cuticular pits on the antennal surface. The s. coeloconica are of two types: one type contains one to three sensory neurons with double sensillar walls penetrated by slit-like pores, whereas the second type contains four sensory neurons with non-porous double sensillar walls. The s. chaetica have a flexible socket and a thick non-porous sensillum wall and contain four sensory neurons that send unbranched dendrites to a terminal pore. A fifth sensory neuron of the s. chaetica terminates in a tubular body at the base of the hair. S. basiconica and coeloconica are normally distributed over the entire antennal flagellum, with a concentration in the middle segments; s. trichodea have three areas of concentration on the 5th, 10th and 14th flagellar segments. Sensilla chaetica are most abundant on the terminal segment. Locusts raised in solitary conditions have more olfactory sensilla (s. basiconica and s. coeloconica) than crowd-reared locusts. The difference in sensillar numbers is more evident in adults than in nymphs. These results suggest that differences in the odour-mediated behaviour of nymphs and adults, and between the phases of S. gregaria, may be attributable to differences at the sensory input level.     

Sex-specific antennal sensory system in the ant <Emphasis Type="Italic">Camponotus japonicus</Emphasis>: structure and distribution of sensilla on the flagellum

The antennae are a critically important component of the ant’s highly elaborated chemical communication systems. However, our understanding of the organization of the sensory systems on the antennae of ants, from peripheral receptors to central and output systems, is poorly understood. Consequently, we have used scanning electron and confocal laser microscopy to create virtually complete maps of the structure, numbers of sensory neurons, and distribution patterns of all types of external sensilla on the antennal flagellum of all types of colony members of the carpenter ant Camponotus japonicus. Based on the outer cuticular structures, the sensilla have been classified into seven types: coelocapitular, coeloconic, ampullaceal, basiconic, trichoid-I, trichoid-II, and chaetic sensilla. Retrograde staining of antennal nerves has enabled us to count the number of sensory neurons housed in the different types of sensilla: three in a coelocapitular sensillum, three in a coeloconic sensillum, one in an ampullaceal sensillum, over 130 in a basiconic sensillum, 50–60 in a trichoid-I sensillum, and 8–9 in a trichoid-II sensillum. The basiconic sensilla, which are cuticular hydrocarbon-receptive in the ant, are present in workers and unmated queens but absent in males. Coelocapitular sensilla (putatively hygro- and thermoreceptive) have been newly identified in this study. Coelocapitular, coeloconic, and ampullaceal sensilla form clusters and show biased distributions on flagellar segments of antennae in all colony members. The total numbers of sensilla per flagellum are about 9000 in unmated queens, 7500 in workers, and 6000 in males. This is the first report presenting comprehensive sensillar maps of antennae in ants.     

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     

Mechanoreception by cuticular sensilla on the pectines of the scorpion <Emphasis Type="Italic">Pandinus cavimanus</Emphasis>

On the pectines of scorpions, several types of cuticular receptors are located. Of these receptors, only the chemo- and mechanosensory peg sensilla have been studied so far while the response characteristics of the long, straight hair sensilla are unknown. As these sensilla protrude in the walking direction and to the ground, we assume that these receptors are most likely involved in observed reflex behaviours. The sensilla constitute rather robust shafts, comparable to other touch-receptors. Their innervation pattern reveals that 5-6 sensory cells are associated with one sensillum. It was possible to record up to three different spike classes (units) which could be distinguished by size, response characteristics and conduction velocity. Two units were analysed in more detail. The response characteristics showed two phasic units, one large and one small, coding the velocity of a stimulus. One medium-sized unit showed phasic-tonic characteristics, coding also the duration of a stimulus. Taking together the morphological and electrophysiological results, we suggest that these sensilla belong to the group of long hair sensilla distributed all over the scorpion body. Furthermore, their response characteristics and the timing between sensory and motor activity within the pectine nerve enable them to be involved in reflex behaviours.