The role of pore structures in the selective permeability of antennal sensilla of the desert burrowing cockroach, Arenivaga sp

To obtain information about the chemical composition of pore structures in antennal sensilla, the antennae were exposed to lipid solvents, or they were prepared to show negative-contrast images in electron micrographs. A heavy-metal tracer, lanthanum nitrate, was also used to indicate the permeability of the receptors to water. The grooves of the large grooved peg open into tubular cavities containing electronopaque material, through which stimulatory molecules must pass to reach the sensory dendrites at the center of the sensillum. The material in these cavities was removed by chloroform or acetone, suggesting a lipid composition. Lanthanum penetrated this receptor only after it had been exposed to acetone or chloroform. Strands at pores of the thin-walled pegs were also removed by the lipid solvents, and the water-soluble tracer failed to penetrate these receptors unless they had been previously exposed to chloroform or acetone. The pore structures appear to be hydrophobic, allowing entry of lipid-soluble substances, while preventing passage of water. The differential action of the solvents on the various types of sensilla suggests that receptor discrimination among different classes of chemical stimuli may be partially determined by the chemical properties of structures at the sensillar pores.     

Antennal chemoreceptors of the desert burrowing cockroach, Arenivaga sp

The antennae of Arenivaga have six types of chemoreceptor sensilla. Some of these have unusual morphological features which may be adaptations for survival in a dry habitat. The sensory dendrites are well protected by cuticular structures, and in some receptors stimulatory molecules must pass through long channels or through pores filled with strands to reach the sensory cells. Large grooved pegs (possibly pheromone receptors) are numerous on antennae of adult males, and grooved sensilla are described here in detail for the first time. Thin-walled pegs, present in males and females, do not have pore tubules or hollow filaments as observed in many other insects. Rather, they contain structures designated here as pore strands, since they have a dense core rather than a light center as previously described for pore tubules and filaments. These strands do not appear to be evaginations of the dendritec membrane, but are probably formed in association with the cuticular structures of the sensilla.     

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

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

Ultrastructure of the galeal sensory complex in adults of the Colorado potato beetle, Leptinotarsa decemlineata

ABSTRACT. The structure of galeal sensilla of the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae), is described using electron microscopical methods. Previous electro-physiological studies indicate that these sensilla respond to amino acids, sucrose and plant saps. One physiological type is particularly sensitive to L-alanine and gamma amino butyric acid (GABA).
Three morphologically different types of sensilla occur on the galeal tip. The more numerous apical pegs are not distinguishable from one another on the basis of external structure, although they differ physiologically. Five sensory cells are associated with most apical pegs. One apical peg, the α-sensillum, contains only four cells. All apical pegs have one cell with a tubular body. The remaining cells have unbranched dendrites and are associated with a single apical pore.
Apical hairs differ from the apical pegs by having double innervation. Within the hair shaft, a dendritic sheath is lacking and the sensillar sinus extends to the base of the hair. The function of this hair type is not known.
Numerous mechanosensory hairs which surround the other sensilla are singly innervated and contain a tubular body at the level of the outer dendritic segments.     

Response properties of chemosensory peg sensilla on the pectines of scorpions

By behavioral and anatomical criteria, the pectinal sensory appendages of scorpions appear to be chemoreceptive organs specialized for detection of substances on substrates. These comb-like, midventral appendages contain tens of thousands of minute (<5 μm), truncated setae, called pegs, arranged in dense, two-dimensional arrays on the ventral surface. In this study we used extracellular recording techniques to examine spontaneous and stimulated activity of sensory neurons within individual pegs. Chronic recordings lasting several days showed long-term fluctuations in spontaneous activity of sensory units in single peg sensilla, with peak activity coinciding with the animal's normal period of foraging. Several units were identified by the stereotypical waveforms of action potentials they elicit. Near-range olfactory stimulation of peg sensilla by volatile alcohols, aldehydes, ketones, esters, and carboxylic acids produced dose-dependent patterns of neural response. Contact stimulation with these chemicals, or water, or mechanical deflection of the peg tip also evoked activity in identifiable units. The peg sensilla appear to be broadly sensitive to odorants and tastants, suggesting they function similarly to the antennae of mandibulate arthropods. Accepted: 12 April 1997     

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

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

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.     

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.     

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 the terminal sensilla on the maxillary palps of Locusta migratoria (L.), and changes associated with moulting

Summary The basic structure of the terminal sensilla of Locusta migratoria resembles that of Schistocerca gregaria. There are commonly six or ten neurons whose dendrites extend almost to the opening of the peg. Proximally the dendrites are clothed by a neurilemma cell which also encloses a basal cavity through which their ciliary region passes. The tormogen cell encloses the receptor-lymph cavity and actively secretes material into it. The receptor-lymph cavity and the basal cavity are quite separate.The development of new pegs at a moult is described. After apolysis the scolopale extends across the subcuticular space and protects the dendrites, which remain in a functional condition until shortly before ecdysis. As the trichogen cell grows out to form a new peg the tip is surrounded by a mass of electron-dense material, probably derived from the receptorlymph cavity. The function of this material is unknown. Regeneration of the dendrites is considered.The possible mechanism by which the tip of the peg opens and closes is considered and the general structure of the organule is discussed in relation to functioning.     

Fine structure of the antennal receptors of the bed bug, Cimex lectularius L.

Sensilla on the antenna of the bed bug, Cimex lectularius, were studied with the scanning and transmission electron microscope. Those which display a tubular body in the dendrite ending are presumed to have a mechanoreceptor function (bristles of type A, flat plate of type B). Bristles of type A1 contain additional dendrites which terminate at the tip of the bristle and may be gustatory receptors. Sensilla with pores in the hair wall are supposed to have an offactory, humidity and/or temperature receptor function (pegs and hairs of types C, D, E). Hairs of type E contain receptors for the alarm pheromones of the bed bug. Special attention has been paid to the pore structures and epicuticular layers of these sensilla. Possible differences in stimulus conduction are discussed between (i) sensilla with a simple wall and pores with pore tubules (types D and E) and (ii) the ribbed pegs (type C), which have a complex wall structure and spoke channels. The immersed cones of type F have a peculiar innervation, which has not been described previously. Two dendrites are held closely together by a third flat dendrite which wraps around them in the region of the outer segment. Coupling structures were found between the central dendrites, and between these and the third enveloping dendrite. Possible functions of this unique innervation are discussed. The dendrites innervating type D are grouped in three to eight bundles by multiple sheaths. The term thecogen cell is introduced to denote the innermost of the three sheath cells of a sensillum (the outer being the tormogen and the trichogen cell) which builds the dendrite sheath during ontogeny. Comparative morphometry revealed type-specific differences in the length and diameter of the dendrites. Some axons were found to lack any glial or perineurial sheath. Microorganisms were observed in the antennal tissue of several animals.     

Scanning electron microscope study of the palar pegs of three species of Corixidae (Hemiptera)

Scanning electron microscope studies were performed on the palar pegs of male forelegs of Trichocorixa sexcincta, Sigara alternata and Hesperocorixa interrupta. The studies showed the pegs to be basically conical and chevron-ridged, with a wide base that tapered of to a narrow apex. Clear species differences were apparent with respect to the size of the peg, the number of ridges on an individual peg, and the number and arrangement of the pegs on the pala. The possible behavioural and evolutionary significance of the palar pegs is discussed in relation to mating.     

The fine structure of the terminal sensilla on the maxillary palps of Schistocerca gregaria (Forskål) (Orthoptera,Acrididae)

Summary The fine structure of the terminal sensilla on the maxillary palps of Schistocerca gregaria has been investigated. Most organules include six neurons with dendrites extending to the tip of the cuticular peg, the opening of which is controlled so that the dendrites are not always exposed. The neurons are isolated from each other by a neurilemma cell and two other glial cells, while typical epidermal cells containing dense bundles of microtubules support the whole group of cells. At the poles of the neurons are specialised areas in which the cytoplasm is differentiated from that elsewhere. It contains a large number of mitochondria and small helical structures, while close to it are characteristic spheres of membranes, termed onion bodies, in various stages of development.It is suggested that the fluid bathing the distal parts of the dendrites and exuding from the tip of the peg has a number of specialised functions. It is probably concerned in forcing open the tip of the peg by hydrostatic pressure, it prevents the exposed tips of the dendrites from desiccating and it acts as a transmitter in which chemicals on the surfaces touched by the sensillum must dissolve before reaching the dendrites. This fluid may be produced by the neurilemma cell or by the neurons themselves. Closure of the pegs does not seem to produce any material reduction in the overall loss of water by the insect.Each neuron sends an axon to the brain; there is no peripheral fusion of axons. Possibly one neuron has a mechanoreceptor function, although no specialised terminal at the base of the peg has been observed. The concentration of mitochondria at either end of the neuron may be concerned in the production of action potentials, while the cavity of the peg and tormogen cell perhaps has a role in the conduction of the receptor potential to the perikaryon. Intercellular connections are such as to give mechanical stability to the cells of the organule and permit transport between the cells. Extracellular tubules extending from the wall of the peg into the cell complex may serve to anchor the peg during the moulting process.We are grateful to the Royal Society for a grant for the purchase of an ultramicrotome, to the Anti-Locust Research Centre for supplying the locusts, and to various colleagues for assistance and advice. We are also indebted to the Cambridge Instrument Company Ltd. for permission to publish Figs. 2 and 3.     

Structure of galeal sensory complex in adults of the red turnip beetle,Entomoscelis americana brown (Coleoptera,Chrysomelidae)

Summary The internal and external structure of the galeae of the adult red turnip beetle, Entomoscelis americana, was studied using SEM and TEM. The galea broadens from base to truncated tip and its sides are of thick, sculpted cuticle invested with pores and coarse spines. The tip is of thinner, flexible cuticle covered with 8–12 uniporous, blunt-tipped apical pegs and a single, aporous, sharply-pointed apical hair.The coarse spines are singly innervated probable mechanosensilla owing to the tubular body at the distal end of the dendrite. These sensilla likely act as tactile hairs monitoring galeal-effected movements of food particles into the functional mouth. The pores are associated with glands within the galea. The function of the presumed secretion is not known but may be to keep objects and dried saliva from sticking to the mouthparts.The apical pegs are innervated by five neurons, each producing a single dendrite. Four dendrites enter the single peg lumen and communicate with the terminal pore. The fifth differentiates into a tubular body that inserts into the peg base. These are typical insect contact chemosensilla that, because of their location, would taste incoming food.The apical hair has no pores but is innervated by two neurons, each extending a dendrite into the hair lumen in chemosensillar fashion. The sensory mode of this sensillum is unknown but is probably not mechanoor chemoreception. Many of its features, reminiscent of taste hairs, lead us to hypothesize that it represents a one-time chemosensillum recently modified to a new form and sensory mode.Because larval and adult E. americana share similar food plant requirements, we hypothesize that similarities will be seen in their mouthpart sensilla. Comparisons of the adults and larvae show the common features between their respective galeal taste hairs are only those of insect contact chemosensilla in general. However, the adult apical hair and the larval medial sensillum show striking specific structural similarities. We propose that these are true structural and functional homologues.     

Ultrastructural diversity in the pore canal systems of amphipod crustaceans

Transmission electron microscopy was used to analyze the structure and organization of the intracuticular pore canal system in 34 species of amphipod crustaceans. Pore canals were detectable in all species, including those that, from scanning electron microscopy, had been considered to lack them. Canal structure ranges from simple transcuticular passages of uniform diameter to more elaborate systems with distal canal dilatations variously equipped with electron dense collars, tubular filaments, single or multiple channels leading to the surface and transverse partitions separating canal contents of different electron densities. Considerable branching between tubular elements of the canal system is evident in many species. In most species the canals communicate with the outsidevia epicuticular channels. Even in those species in which this communication was not established, their epicuticle generally contains abundant cavities with external pores. Although some consistency in structure and organization is present among a few groups of taxonomically related species, pore canal characteristics generally could not be correlated with habitat or life style. It is suggested that the structural and organizational variety present reflects a considerable array of functions among amphipod pore canal systems.     

Fine structure of antennal sensilla coeloconica of culicine mosquitoes

The sensilla coeloconica (pegs in pits) previously mis-identified as campaniform organs, at the tip of the antennae of female Aedes aegypti L. and Culexpipiens (L.) are described. Each sensillum is innervated by three bipolar neurons: the dendrites of two are unbranched whereas the distal portion of the third is folded into tightly packed lamellae. One unbranched dendrite extends to the tip of the peg and the other ends near the base of the peg. The lamellae-bearing dendrite terminates 4-5 μ beneath the base of the peg. Chemo- and thermoreception are the proposed functions for the sensillum.     

Mapping and ultrastructure of antennal chemosensilla of the wheat bug Eurygaster maura

Antennae of the wheat stink bug Eurygaster maura L. (Hemiptera: Scutelleridae) were investigated to elucidate structure and distribution of antennal chemosensilla in females. Five type of sensilla were identified and characterized using a scanning electron microscope (SEM) and a transmission electron microscope (TEM). Type 1 sensilla are mechanical and contact chemoreceptors with a single apical pore. Types 2 and 3 sensilla are multiporous chemoreceptors both with typical features of olfactory sensilla. Type 4 are multiporous peg-like sensilla, short and with a grooved surface. Type 5 are sensilla coeloconica with a smooth and aporous peg completely inserted in a sub-cuticular chamber. All types are distributed on the two flagellar segments, but we considered only the apical flagellomere in which the largest number of sensilla are located. The most abundant sensilla are type 3, while the less numerous are type 5. All types, except type 2, decreased in number from the tip to the base of the segment. The lower density of sensilla was recorded on the dorsal-internal part of the apical antennomere, while the higher density was recorded on the opposite side (external-ventral).     

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)     

Functional anatomy of sensory structures on the antennae of Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae)

The ultrastructure and distribution of sensilla on the antennae of the cabbage stem flea beetle, Psylliodes chrysocephala, were investigated using scanning and transmission electron microscopy techniques. Eight different sensillar types were distinguished. These were; hair plate sensilla, sensilla chaetica, three types of sensilla trichodea, sensilla basiconica, grooved peg sensilla and styloconic sensilla. The sensilla chaetica are known to be gustatory receptors. Ultrastructure indicates that the hair plate sensilla and sensilla trichodea type one are probably mechanoreceptors, whilst the sensilla styloconica are probably thermo-hygro receptors. These thermo-hygroreceptors are unusual in that they are innervated by two sensory cells (one hygroreceptor and one thermoreceptor) rather than the more usual triad. The remaining four sensillar types all have a porous hair shaft, indicating an olfactory role. One of these (the grooved peg sensillum) may also have a thermoreceptive function. No sexual dimorphism was found in the structure, number or distribution of the antennal sensilla.     

Sensilla basiconica (grooved pegs) on the antennae of female mosquitoes: electrophysiology and morphology

The sensilla basiconica or grooved pegs of mosquitoes consist of two morphologically and functionally distinct classes. Short grooved pegs house sensory cells that are excited by lactic acid whereas lactic acid-excited cells are absent in long grooved pegs. This correlation was found in four species of mosquito examined:Aedes aegypti, Aedes atropalpus, Aedes epactius, andCulex pipiens. InAedes epactius, both short and long grooved pegs house receptors that are sensitive to butyric acid.