Sensilla and Cuticular Appendages on the Female Abdomen of Lasioptera rubi (Schrank) (Diptera,Cecidomyiidae)

Studies by SEM and TEM revealed 6 types of integumental appendages on female uromeres VIII-X in Lasioptera rubi: microtrichia, not innervated; spines, probably without sensory function; nonporous sensory hairs, each containing one dendrite ending with a tubular body indicating a tactile function; uniporous sensory hairs, each innervated partly by 3 dendrites indicating a chemosensory function, partly by an additional dendrite with a tubular body indicating a tactile function; scoop-like sensilla, each containing partly a branched structure of dendrites in the distal half of the sensillum indicating an olfactory function, partly an unbranched dendrite ending at a pore near the base of the sensillum, most probably registrating chemical stimuli by contact or gustation; finally, nonporous bristles, all or some of them innervated, in a manner indicating a tactile function. In addition, two scolopophorous proprioceptors were found inside uromere X. The nonporous sensory hairs, the uniporous sensory hairs and the scolopophores may be used by the midge to determine the mechanical and chemical properties of potential oviposition sites. The spines and nonporous bristles may function as conidia carriers.     

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.     

The gustatory sensilla on the endophytic ovipositor of Odonata

The present paper aims at describing the fine structure of coeloconic sensilla located on the cutting valves of the endophytic ovipositor of two Odonata species, the anisopteran Aeshna cyanea (Aeshnidae) and the zygopteran Ischnura elegans (Coenagrionidae), by carrying out parallel investigations under SEM and TEM. In both species these coeloconic sensilla are innervated by four unbranched neurons forming four outer dendritic segments enveloped by the dendrite sheath. One dendrite terminates at the base of the peg forming a well developed tubular body, while the other three enter the peg after interruption of the dendrite sheath. The cuticle of the peg shows an apical pore and a joint membrane. This last feature, together with the tubular body and the suspension fibers, represent the mechanosensory components of the sensillum while the pore and the dendrites entering the peg allow chemoreception. The ultrastructural organization of these coeloconic sensilla is in agreement with the one reported for insect gustatory sensilla. Our investigation describes for the first time typical insect gustatory sensilla in Odonata. Electrophysiological and behavioral studies are needed to verify the role that these structures can perform in sensing the egg-laying substrata.     

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.     

Fine structure of tarsal sensilla of Aedes aegypti (L.) (Diptera: Culicidae)

The tarsi of all three pairs of legs of both sexes of Aedes aegypti (L.) bear spine sensilla, five types of hair sensilla, which are designated A, B, C1, C2 and C3, and campaniform sensilla. Type A and B hairs, spines, and cam-paniform sensilla are innervated by one neuron with a tubular body, a characteristic of cuticular mechanoreceptors. In particular the hairs and spines are tactile receptors and the campaniform sensilla are proprioceptors. The C1, C2, and C3 hair sensilla have the morphological features of contact chemoreceptors. Type C1 and C3 hairs are innervated by five and four neurons, respectively, which extend to the tip of the hair. Type C2 is innervated by five neurons, one of which terminates at the base of the hair in a tubular body while the remaining four extend to the tip of the hair. The role of the type C hairs in oviposition behavior, nectar feeding, and recognition of conspecific females is discussed. Presumed efferent neurosecretory fibers occur near the spine and hair sensilla.     

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.     

The cercal sensilla of the blowfly Lucilia cuprina. I. Structure of the sockets and distal dendritic regions

The functions of the gustatory, olfactory, touch and stress receptors on the cerci of Lucilia cuprina Wied. (Diptera: Calliphoridae) are apparent from the morphology of their distal dendritic segments and associated cuticular structures. Each trichoid mechanoreceptor has a dendrite containing a tubular body at the base of the hairshaft. The suspension fibres and socket septum may be involved in transmitting a stimulus to the dendrite terminal and restoring the hair to its resting position. The campaniform sensilla are considered as trichoid mechanoreceptors with reduced hair shafts and socket structures, reflected in fusion of the suspension fibres into the inner cuticle of the dome and loss of the socket septum. Fusion and reduction of the socket structures is also apparent at the bases of the olfactory pegs. They differ from typical antennal olfactory sensilla in having a flexible socket and relatively thick walls; features which may protect them from damage during ovipositor probing of potential oviposition substrates. The two types of cereal gustatory sensilla differ in their complement of chemosensory dendrites, one has three, the other four, the latter type also has a mechanoreceptive dendrite at the base of the hair shaft. Both types have socket structures resembling those of the trichoid mechanoreceptors.     

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.     

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.     

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.     

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.     

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.     

Antennal contact chemosensilla in Psylliodes chrysocephala responding to cruciferous allelochemicals

Sensilla chaetica, which protrude above all other sensilla on the antenna of Psylliodes chrysocephala L., the cabbage stem flea beetle, were investigated ultrastructurally and found to be innervated by five to six sensory neurones. A dendrite from one of these neurones terminates in a tubular body at the shaft base, whereas dendrites from the others run unbranched to a pore at the shaft tip. Such a structure typifies a sensillum with a combined gustatory/mechanosensory function. Electrophysiological recordings using the tip-recording technique confirmed that this sensillum contains one mechanosensory cell and several chemosensory cells. The chemosensory cells were responsive to host plant chemicals. Sensilla chaetica were also found to be responsive to glucosinolates. One of the sensilla chaetica emerges from a domed area of cuticle on antennomere six. This was found to be relatively less responsive to the chemical stimuli tested and more responsive to mechanical stimulation. It is suggested that the sensilla chaetica are contact chemosensilla, that respond to chemicals present in plant surface waxes when P. chrysocephala antennates a leaf.     

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.     

The cercal sensilla of the blowfly Lucilia cuprina. II. Structure of the enveloping cells and the basal regions of the sensory dendrites

The gustatory, olfactory, touch and stress receptors on the cerci of Lucilia cuprina Wied. (Diptera: Calliphoridae) have either two or three enveloping cells. The gustatory and olfactory sensilla have three enveloping cells: a tormogen, trichogen and thecogen cell. The tormogen and trichogen cells contribute to a sub-cuticular sensillar lumen which divides into two lobes basally. The thecogen cell forms a lumen around the dendrites. Distally the dendrites lie in the contents of the thecogen lumen within the dendritic sheath. Proximally the dendrites embed in the thecogen cell which has an expanded, microlamellate lumen basally. The sensillar lumen of the mechanosensory (trichoid mechanoreceptors and campaniform) sensilla is formed by a single enveloping cell: the presumptive tormogen cell. In trichoid mechanoreceptors the thecogen lumen is restricted to the region of the transitional region of the dendrite whereas the thecogen lumen of campaniform sensilla extends proximally although it is not as well-developed as that of the chemoreceptive sensilla. The dendrites of all sensillum types on the cerci have a granular body in the transitional region: a situation which has not been previously reported in chemoreceptive sensilla although common in the mechanoreceptors of Calliphoridae and Sarcophagidae.     

Antennal sensory organs in the millipede Eurypauropus ornatus: Fine structure of the flagella and globulus

On the antennal tip of Eurypauropus ornatus are 3 threadlike sensilla—the flagella, and a single spheroid sensillum—the globulus. Each of the 3 flagella is innervated by 2 groups of sensory cells. One group contains 4 cells, the other, 5. All cells of the “four group” and 3 of the “five group” are comprised of single cilia and unbranched dendrites which extend along the lumen of the flagellum. Two cells of the “five group” have double cilia and pairs of unbranched dendrites. One pair also enters the flagellum and the other pair terminates beneath the flagellar base to form a concentric array of lamellae. No pores are present in the cuticular wall. Eight sensory cells innervate the globulus. They are arranged in 3 groups, one triplet and 2 pairs, in addition to a single cell. The single cell contains a pair of cilia whose unbranched dendrites differentiate into tubular bodies that are inserted into the base of the globulus. Each of the other 7 sensory cells has a single cilium. Their unbranched dendrites penetrate into the globulus in 3 groups as described for the sensory cells. The dendrites in each group terminate in an individual pore channel at the globulus tip and completely fuse with the electron-dense material that plugs the pore channel. Based on structural similarities to sensilla having known functions, it is probable that the flagella and the globulus are chemoreceptors, the former responding to odors, the latter sensitive to substances in aqueous solution.     

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.     

Morphology and ultrastructure of sense organs on the ovipositor of Trybliographa rapae,a parasitoid of the cabbage root fly

The ovipositor of the parasitoid wasp Trybliographa rapae was examined by scanning and transmission electron microscopy. Characteristic peg-like sensilla with a cuticular ring at the base are found at the tip of the ventral valves, where they occur in a characteristic arrangement of triplets. The unusual basal structure probably protects the sensilla against damage during movement through the substrate and piercing of the host cuticle. These sensilla are each innervated by six dendrites, some of which have lamellated tips, generally considered to be characteristic of thermosensitivity. It is suggested that the remaining dendrites are gustatory, and as such probably respond to factors present in host haemolymph. A second type of peg-like sensillum is found on both the dorsal and the ventral valves. These are set in deep pits so that only the tip of the peg protrudes above the surface of the cuticle. These occur along the length of the ovipositor shaft and ultrastructural studies reveal the pegs to be innervated by a single mechanosensitive dendrite, probably monitoring the movement of the ovipositor through the substrate.     

Gustatory organs of Drosophila melanogaster: fine structure and expression of the putative odorant-binding protein PBPRP2

In Drosophila, as in most insects, gustation is mediated by sensory hairs located on the external and internal parts of the proboscis and on the legs and wings. We describe in detail the organization and ultrastructure of the gustatory sensilla on the labellum and legs and the distribution of PBPRP2, a putative odorant-binding protein, in the gustatory organs of Drosophila. The labellum carries two kinds of sensilla: taste bristles and taste pegs. The former have the typical morphology of gustatory sensilla and can be further subdivided into three morphological subtypes, each with a stereotyped distribution and innervation. Taste pegs have a unique morphology and are innervated by two receptor cells: one mechanoreceptor and the other a putative chemoreceptor cell. PBPRP2 is abundantly expressed in all adult gustatory organs on labellum, legs, and wings and in the internal taste organs on the proboscis. In contrast to olfactory organs, where PBPRP2 is expressed in the epidermis, this protein is absent from the epidermis of labial palps and legs. In the taste bristles of the labellum and legs, PBPRP2 is localized in the crescent-shaped lumen of the sensilla, and not in the lumen where the dendrites of the gustatory neurons are found, making a function in stimulus transport unlikely in these sensilla. In contrast, PBPRP2 in peg sensilla is expressed in the inner sensillum-lymph cavity and is in contact with the dendrites. Thus, PBPRP2 could be involved as a carrier for hydrophobic ligands, e.g., bitter tastants, in these sensilla.