The sensory systems and feeding behavior of leafhoppers. I. The aster leafhopper,Macrosteles fascifrons stål (homoptera,cicadellidae)

The ultrastructure of the sensilla, and other structures, within the stylets and precibarium of Macrosteles fascifrons were examined by transmission and scanning electron microscopy. Precibarium is a new term, defined here, for the canal that precedes the cibarium inside the leafhopper head. Within the precibarium are found 20 chemosensilla and a previously undescribed structure, the precibarial valve. Twelve mechanosensilla, three in each stylet, are found within the maxillary and mandibular stylets. The relationship between all of these structures and feeding by the insect is detailed in a feeding mechanism hypothesis. It is concluded that leafhoppers (and probably all homopterans) utilize the precibarial chemosensilla alone for gustatory discrimination, the stylet sensilla for proprioception, and the precibarial valve for regulation of fluid uptake and compartmentalization of the sensilla.     

Precibarial and cibarial chemosensilla in the whitefly,Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae)

The internal anatomy of the anterior alimentary canal of the whitefly, Bemisia tabaci (Gennadius) (Homoptera: Aleyrodidae) B-biotype, was examined by light, scanning, and transmission electron microscopy to elucidate the location and number of precibarial and cibarial gustatory sensilla. Elucidation of the epipharyngeal organ complex within the precibarium revealed 10 precibarial sensilla located proximal to where the paired maxillary stylets diverge on their retraction. The sensory organ complex within the cibarium consists of 8 sensilla, 6 on the epipharyngeal sclerite with 2 found within the hypopharyngeal sclerite. Fine structure investigation revealed the individual neurons to terminate at sensillar pores, which allow direct contact with passing fluids, thus supporting a chemosensory function. Ultrastructure of the neurons is similar to that of precibarial and cibarial gustatory chemosensilla found in other piercing-sucking insects. Their importance to whitefly feeding is discussed.     

Comparison of antennal sensilla of Anaphes victus and A. listronoti (Hymenoptera, Mymaridae), egg parasitoids of Curculionidae

Antennal sensilla were compared in females and males of two sympatric mymarid Hymenoptera, Anaphes victus and A. listronoti which are, respectively, solitary and gregarious parasitoids of eggs of the carrot weevil Listronotus oregonensis (Coleoptera, Curculionidae). Both species are morphologically very similar in the area where they are sympatric. The external morphology of the sensilla was studied using scanning electron microscopy. Female antennae have seven different types of sensilla, morphologically similar in the two species: trichoid sensilla, which are putative mechanosensilla, sensilla chaetica types 1, 3 and 4, which are presumably contact chemosensilla, and sensilla chaetica type 2 and basiconic and placoid sensilla, which are presumed to be olfactory sensilla. The major difference between the two species is the number of sensilla chaetica type 4, of which 6–9 are found on the antennal club in A. victus, while 10–12 are present in A. listronoti. The antennae of the males of both species are similar in morphology and in the number and distribution of their four types of sensilla, i.e. trichoid sensilla, sensilla chaetica type 1 and basiconic and placoid sensilla. Accepted: 23 November 1998     

Antennal sensilla in five Psychodini moth flies (Diptera: Psychodidae: Psychodinae) pollinators of Arum spp. (Araceae)

The pollination of the genus Arum (Araceae) is mainly achieved by deception, the floral odour mimicking the pollinator ovipositing site. In order to discover the sensory organs involved in this attraction, we have studied the antennae of five species of psychodine moth-flies (former Psychoda sensu lato=Psychodini), pollinators of Arum spp. The antennae of the five Psychodini reveal seven types of sensilla: multiporous tribranched sensilla basiconica (sensilla ascoidea), multiporous sensilla basiconica, multiporous sensilla coeloconica, multiporous sensilla auricillica, uniporous sensilla basiconica, aporous sensilla chaetica, aporous Böhm’s sensilla. Each species possesses three, five or six of these sensillum types. All the multiporous sensilla are probably olfactory receptors while the uniporous sensilla basiconica must possess a contact chemoreceptive function. The multiporous tribranched sensilla basiconica (s. ascoidea), present in all the species, are the best candidates for the reception of the odours given off by the ovipositing sites and the inflorescences of Arum. The multiporous sensilla basiconica and the multiporous sensilla coeloconica may be involved respectively as CO₂ receptors or thermoreceptors. Psychoda phalaenoides, which is the main pollinator of A. maculatum, is the species which possesses the largest number of antennal sensilla. The sexual dimorphism, studied only in Psycha grisescens, as concerns the number of sensilla and the absence of a sensillum type which differ according to sex, is difficult to interpret.     

Variation in antennal sensilla of some hepialid moths; a scanning electron microscope study

Sensilla on the antennae of 5 species of hepialid moths—Aenetus virescens (Hepialinae) and Wiseana cervinata, W. signata, W. umbraculata, and Trioxycanus enysii (Oxycaninae)—were examined with the scanning electron microscope. Only 4 of the 6 types of sensilla previously described from noctuid moths were observed. The arrays of sensilla on oxycanine species are not uniformly distributed, especially on males; most of the sensors on any segment are on the ‘ridge’ of the pectinations, with a sparse distribution of relatively widely spaced sensors on the rest of the cuticle. In all species examined the antennae were sexually dimorphic; this was most apparent in the differing sensilla arrays of Hepialinae and in the difference in size of the antennal pectinations of Oxycaninae.     

Anterior alimentary canal of the pear Psylla,Psylla pyricola foerster (Homoptera,Psyllidae)

Scanning and light microscopy investigations of the anterior alimentary canal of the pear psylla, Psylla pyricola Foerster (Homoptera: Psyllidae), revealed the morphology of the labium and stylets, as well as the presence of sensory structures and a valve in the precibarium. The labium consists of three telescoping segments with an internal labial groove, which surrounds and supports the stylet bundle. Also a part of the labial groove is the internal labial clamp. The stylet bundle is comprised of paired styliform mandibles and maxillae, which interlock to form the food and salivary canals. The stylet bundle proximal to the labium forms a large loop within a membranous crumena. When fully retracted the coiled stylets are under tension. Stylet extension generates increasing tension so that when retracted the stylets readily recoil within the crumena. Penetration of leaf tissues by the stylet bundle is dependent on the interaction between stylet muscles, opening and closing of the labial clamp, the barbed stylet tips, and the ventral position of the labium. Proximal to the crumena the paired stylets separate and diverge at the entrance of the precibarium, which is formed by the interlocking of the epi-and hypopharynges. There are 18 sensory structures in the precibarium, as well as a precibarial valve. These structures appear to be homologous to similar structures observed previously in aphids and leafhoppers. The morphology and the location of the precibarial sensilla suggest that, like the precibarial sensory organs of aphids and leafhoppers, they are gustatory and probably mediate acceptance or rejection of plant fluids, thus playing a major role in locating tissues for feeding.     

Ultrastructure of sensilla on the maxillary and labial palps of three species (Coleoptera: Staphylinidae: Staphylininae)

Most species of Staphylinidae are predators in an agroecosystem. They acquire prey information from their environment through receptors. In this study, the sensilla on maxillary and labial palps of Philonthus kailiensis, Philonthus lewisius and Quedius robustus were examined with scanning electron microscopy to identify and analyse the external morphology and distribution of the sensilla to enhance our knowledge of the sensilla of Staphylinidae and provide a rationale of taxonomical studies on the two genus. Results showed that the sensilla are classified into six types: Böhm bristles, sensilla chaetica, sensilla furcate, sensilla coeloconica, sensilla placodea and sensilla basiconica. No sexual dimorphism exists among the three species. The relationships and functions of sensilla on maxillary and labial palps were also speculated. There may be a certain correlation between the sensilla on maxillary and labial palps of the staphylinid and its habitat.     

Ultrastructure and distribution of sensilla on the antennae of female fig wasp Eupristina sp. (Hymenoptera: Agaonidae)

In the species‐specific and obligate mutualism between the fig (Moraceae: Ficus spp.) and its pollinator (Hymenoptera: Agaonidae), the continuity of lifecycle of both partners completely depends on the female pollinator's ability to detect receptive figs. To better understand the chemical location mechanism, we examined the antennae and their sensilla of the female fig pollinator Eupristina sp. using scanning electron microscopy (SEM) and transmission electron microscopy (TEM). The antennae of female Eupristina sp. are geniculated, and in total, there were seven types of sensilla found on the antennae: two types of multiporous placoid sensilla (type 1 is sausage‐like and type 2 is rounded), sensilla trichodea (ST), basiconic sensilla (BS), chaetica sensilla (ChS), coeloconic sensilla (CoS), and one specialized sensillum classified as sensillum obscurum (SO). We described external morphology, abundance, distribution, ultrastructure and discussed putative functions. We inferred from their ultrastructures as chemoreceptors that two types of multiporous placoid sensilla, BS and CoS, were innervated by sensory neurons. The aporous type ST, ChS, and SO were not innervated by dendrites which may function as mechanoreceptor/proprioceptor. These results were also discussed in relation to the interaction between Eupristina sp. and its host fig.     

Ultrastructure and distribution of sensilla on the maxillary and labial palps of Chlorophorus caragana (Coleoptera: Cerambycidae)

Chlorophorus caragana is a species of long‐horned beetle that damages Caragana davazamcii Sancz. (Fabales: Papilionaceae) bushes in desert areas in China. The beetles cause substantial damage to local forestry plantations and the environment. Sensilla on the maxillary and labial palps of coleopterans a allow the insects to recognize their host plants. We used scanning and transmission electron microscopy to study the ultrastructure, distribution, and abundance of various sensilla on the maxillary and labial palps of C. caragana. We found four types of sensilla including ten subtypes: one of Böhm's bristles, three of sensilla chaetica, one of digitiform sensilla, and five of sensilla twig basiconica. The types and distribution of the sensilla on the maxillary and labial palps were highly similar between males and females. Finally, this article discusses the functions of the sensilla of related species in recognizing hosts and the significance of gustation studies in the context of the control of C. caragana.     

Comparative external ultrastructure and diffusion pathways in styloconic sensilla on the maxillary galea of larval Mamestra configurata (Walker) (Lepidoptera: Noctuidae) and five other species

The external ultrastructure of sensilla on the maxillary galea are investigated in Mamestra configurata and five other lepidopterous larvae using scanning electron microscopy. The galea and lacinia, comprising the mesal lobe of the maxilla, are either completely separate, fused, or incompletely fused in these species. The distal surface of the mesal lobe of all species examined bears two styloconic sensilla, three basiconic sensilla, and three trichoid sensilla, whereas the midventral wall of this lobe bears a campaniform sensillum. The latter sensillum is visible in only three of the six species examined. The styloconic and basiconic sensilla occupy a ventro anterior location, whereas the trichoid sensilla are positioned dorsoposteriorly. Interspecific comparisons of galeal size, as well as sensillar size, shape, and position are made for all species. The styloconic sensilla are the only sensillar type permeable to an aqueous solution of cobalt chloride when viewed by brightfield light microscopy in all species examined. Cobalt ions permeate through the terminal pore of each styloconic peg and percolate through the fenestrated fibrillar pore matrix, located directly below the pore. These ions permeate along the dendritic channel and accumulate in the adjacent sensillar sinus surrounding the peg and/or style by way of a presumably permeable dendritic sheath in all species, but to varying extents. The cuticular sidewall pores surrounding the terminal pore also appear to be permeable to cobalt ions in all the species examined. In most species examined, the styloconic sensilla are only minimally permeable to mercury ions. In these species, mercury ions permeate through the terminal pore, but become trapped within the plug of fenestrated fibrils within it. The sidewall pores are not permeable to mercury ions in any of the species examined. The styloconic sensilla are not permeable to lead ions in M. configurata or Malacosoma lutescens, the only species tested. © 1996 Wiley-Liss, Inc.     

Food selection in adults and larvae of three species of Lepidoptera: a behavioural and electrophysiological study

Adults and larvae of Spodoptera littoralis, Heliothis virescens and Heliothis armigera were tested with a range of sugars, amino acids, sugar alcohols and allelochemics. Feeding behaviour was correlated with the electrophysiological responses of maxillary styloconic sensilla in the larvae and proboscis styloconic sensilla in the adult. The neural response was more vigorous in larvae than in adults but otherwise the spectra of responses were similar in the two life stages. Phagostimulants and antifeedants stimulated maximally different sensilla in larvae but not in adults. The responses of adult sensilla to sugars and amino acids were significantly correlated to behaviour in all three species, but only in H. armigera was there a similar correlation with the sugar alcohols and allelochemics.     

Sensilla on antennae,ovipositor and tarsi of the larval parasitoids,Cotesia sesamiae (Cameron 1906) and Cotesia flavipes Cameron 1891 (Hymenoptera: Braconidae): a comparative scanning electron microscopy study

Two braconid parasitoids of cereal stemborers in eastern Africa, Cotesia sesamiae and Cotesia flavipes, have been shown to display a similar hierarchy of behavioural events during host recognition and acceptance. In order to understand the mechanisms underlying host recognition and acceptance, the morphology of antennal sensilla on the last antennomeres, on the ovipositor, and on the fifth tarsomere and pretarsus of the prothoracic legs tarsi were studied using scanning electron microscopy followed by selective silver nitrate staining. It appeared that female C. sesamiae and C. flavipes shared the same types and distribution of sensory receptors, which enable them to detect volatiles and contact chemical stimuli from their hosts. In both parasitoids, four types of sensilla were identified on the three terminal antennomeres: (i) non-porous sensilla trichodea likely to be involved in mechanoreception, (ii) uniporous sensilla chaetica with porous tips that have gustatory functions, (iii) multiporous sensilla placodea, which are likely to have olfactory function, and (iv) sensilla coeloconica known to have thermo-hygroreceptive function. The tarsi of both parasitoids possessed a few uniporous sensilla chaetica with porous tips, which may have gustatory functions. The distal end of the ovipositor bore numerous dome-shaped sensilla. However, there were no sensilla coeloconica or styloconica, known to have gustatory function in other parasitoid species, on the ovipositors of the two braconid wasps.     

Ultrastructure of the antennal sensilla of the cockroach-egg parasitoid,Tetrastichus hagenowii (Hymenoptera: Eulophidae)

Eggs of a number of cockroach species are parasitized by Tetrastichus hagenowii. The ultrastructure of the sensilla on the antennae of females and males was examined by scanning and transmission electron microscopy. The females have two types of multiporous plate sensilla while the males have only one. Type 1 is found in females and males and has a relatively thin cuticular wall and many pores, while type 2 is found only in females and has a relatively thick cuticular wall and few pores. Both sexes have nonporous, thick-walled, socketed hairs; multiporous, nonsocketed hairs; multiporous, thick-walled pegs; and terminal hairs. In addition, males have multiporous, nonsocketed, long hairs. The sensilla are similar, in many respects, to the sensilla of other chalcid parasitoids. The antennal sensilla of female T. hagenowii are probably involved in ovipositional behavior. The multiporous, long hairs of the male possibly receive stimuli during mating behavior A chemoreceptive function is proposed for the multiporous plate sensilla.     

Comparative study of the antennal sensilla of five species of root maggots: Delia radicum L., D. floralis F., D. antiqua Mg., D. platura MG. (Diptera : Anthomyiidae) and Psila rosae F. (Diptera : Psilidae)

A comparative study of the antennal sensilla of Delia radicum L., D. floralis F., D. antiqua Mg., D. platura Mg. (Diptera : Anthomyiidae) and Psila rosae F. Diptera Psilidae) is undertaken. For both sexes of each species, the type, distribution, and density of sensilla are determined. All 5 species have trichoid (olfactory) and grooved (olfactory) sensilla. Basiconica I (blunt) sensilla (olfactory) are found on each of the species examined, except D. platura. Basiconica II (tapered) (olfactory) and clavate (olfactory) sensilla are found only on Delia species. Also, only Delia species have single-chambered, dorsal pits, and these contain basiconic II pit sensilla (olfactory). Common to all 5 species is a multi-chambered ventral pit (olfactory). In the ventral pit, all 5 species have grooved pit sensilla (olfactory). In addition to this type of sensillum the Delia species have smooth-walled conical pit sensilla (hygro-/thermosensitive) and P. rosae has granular pit sensilla (hygro-/thermosensitive). Smooth-walled tapered pit sensilla (hygro-/thermosensitive) are found in D. radicum. Similarities and differences in the density of surface sensilla between dorsal and ventral funicular surfaces, male and female flies, and oligophagous (D. antiqua, D. radicum, D. floralis and P. rosae) and polyphagous (D. platura) species are compared. Several differences in sensillum density between the dorsal and ventral funicular surfaces are observed, but these do not fit into a consistent trend. Except for D. radicum, there are differences in sensillum density between male and female flies. For the oligophagous species, females have a greater sensillum density, whilst for the polyphagous D. platura males have a greater sensillum density. Comparisons between species show the greatest differences between the Delia species and P. rosae, and within the 4 Delia species, differences in sensillum density do not correlate with host range or body size.     

The antennae of neopseustid moths: Morphology and phylogenetic implications, with special reference to the sensilla (Insecta, Lepidoptera, Neopseustidae)

The antennae of Lepidoptera Neopseustidae were examined with the scanning electron microscope. The studied species, Nematocentropus cfr. omeiensis, Neopseustis meyricki, Synempora andesae, Apoplania valdiviana and Apoplania penai possess nine types of antennal flagellum sensilla: multiporous large sensilla basiconica, multiporous thin sensilla basiconica, multiporous small sensilla basiconica, multiporous sensilla trichodea, multiporous sensilla coeloconica; uniporous sensilla chaetica; aporous sensilla chaetica, aporous stylus-shaped sensilla chaetica, aporous sensilla styloconica.The multiporous sensillum type here termed “multiporous large sensillum basiconicum” is unknown from other Lepidoptera and probably constitutes an autapomorphy of the family Neopseustidae. This sensillum type is remarkable by having a single base in female Apoplania and Synempora while in male Apoplania it has a bifid or trifid base, and in male Synempora it is composed of two or three incompletely separated hairs. This may be the first recorded example of a sexually dimorphic lepidopteran sensillum type. The stylus-shaped sensillum chaeticum is a primitive type which occurs only in some lower Lepidoptera.     

Ultrastructure of chemosensory tarsal tip-pore sensilla of Argiope spp. Audouin, 1826 (Chelicerata: Araneae: Araneidae)

While chemical communication has been investigated intensively in vertebrates and insects, relatively little is known about the sensory world of spiders despite the fact that chemical cues play a key role in natural and sexual selection in this group. In insects, olfaction is performed with wall–pore and gustation with tip-pore sensilla. Since spiders possess tip-pore sensilla only, it is unclear how they accomplish olfaction. We scrutinized the ultrastructure of the trichoid tip-pore sensilla of the orb weaving spider Argiope bruennichi—a common Palearctic species the males of which are known to be attracted by female sex pheromone. We also investigated the congener Argiope blanda. We examined whether the tip-pore sensilla differ in ultrastructure depending on sex and their position on the tarsi of walking legs of which only the distal parts are in contact with the substrate. We hypothesized as yet undetected differences in ultrastructure that suggest gustatory versus olfactory functions. All tarsal tip-pore sensilla of both species exhibit characters typical of contact-chemoreceptors, such as (a) the presence of a pore at the tip of the sensillum shaft, (b) 2–22 uniciliated chemoreceptive cells with elongated and unbranched dendrites reaching up to the tip-pore, (c) two integrated mechanoreceptive cells with short dendrites and large tubular bodies attached to the sensillum shaft's base, and (d) a socket structure with suspension fibres that render the sensillum shaft flexible. The newly found third mechanoreceptive cell attached to the proximal end of the peridendritic shaft cylinder by a small tubular body was likely overlooked in previous studies. The organization of tarsal tip-pore sensilla did not differ depending on the position on the tarsus nor between the sexes. As no wall-pore sensilla were detected, we discuss the probability that a single type of sensillum performs both gustation and olfaction in spiders.     

Antennal morphology of the endogean carabid genus typhlocharis (Coleoptera: Carabidae: Anillini): Description of sensilla and taxonomic implications

The antennal morphology and chaetotaxy were studied in 52 species of the endogean carabid genus Typhlocharis, using scanning electron microscopy and light microscopy. The antennae are composed of 11 antennomeres (scape, pedicel, and nine flagellomeres). We found considerable variation between species in the third antennomere, with short‐stem and long‐stem forms, and flagellomere morphology, distinguishing two morphs: rounded (subovoid, subspheric and subquadrate, morph 1) and reniform shapes (morph 2). Antennal sensilla are grouped in six types of sensilla trichodea, three types of sensilla basiconica, one type of sensilla coeloconica, and one type of sensilla campaniformia. The distribution of sensilla along the antennomeres is described. The “rings” of trichoid sensilla in the antennomere body are affected by its shape and there is interspecific variation in the pattern of sensilla coeloconica in antennomere 11°, a novelty for the genus. The types of sensilla found in Typhlocharis are compared to those described in other Carabidae and the potential functionality and taxonomic interest of those variable antennal features are discussed. A correlation between the flagellomere morphology and the presence/absence of a stridulatory organ is suggested. The study also allowed comparing the observation of antennal features by SEM and light microscopy. J. Morphol., 2013. © 2013 Wiley Periodicals, Inc.     

External morphology of antennal sensilla of four species of adult flea beetles (Coleoptera : chrysomelidae : alticinae)

External morphology of antennal sensilla of both sexes of Phyllotetra cruciferae (Goeze) and Psylloides punctulata Melsh, both of which feed on the mustard family Cruciferae, and Epitrix cucumeris (Harris) and Psylloides affinis (Paykell), both of which feed on the nightshade family Solanaceae, was studied using scanning electron microscopy. All belong to Alticinae (Coleoptera : Chrysomelidae). The number and distribution of antennal sensilla were also determined. Eight types of sensilla could be distinguished on the flagella of the species examined: sensilla chaetica; sensilla trichodea I and II; long sensilla basiconica I and II; and short sensilla basiconica I, II and III. The sensilla chaetica are probable tactile mechanosensilla, whereas both types of sensilla trichodea and long sensilla basiconica likely function in olfaction. Suggested functions for the short sensilla basiconica I, II and III include hygro-/thermoreception and chemoreception.The lack of sexual dimorphism in antennal structure and in types as well as number of sensilla, indicates that the sensilla probably have similar functions in both sexes. Probably most of the antennal olfactory sensilla are involved in host location and recognition. Any correlation between number of a particular type of sensillum or total number of all types and general host preference is not apparent.     

Immunolocalization of pheromone-binding protein and general odorant-binding protein in olfactory sensilla of the silk moths Antheraea and Bombyx

The distribution of odorant-binding proteins among olfactory sensilla of three moth species was studied by immuno-electron microscopy. Two polyclonal antisera were used in a post-embedding labelling protocol on sections of cryo-substituted antennae. The first was directed against the pheromone-binding protein (PBP) of Antheraea polyphemus, the second against the general odorant-binding protein (GOBP) of the same species. Immunoblots showed that these antisera were highly specific; both antisera did, however, cross-react with related proteins in the related species A. pernyi, and in the bombycid moth B. mori. PBP and GOBP were localized only in olfactory sensilla trichodea and sensilla basiconica, the principal site being the sensillum lymph surrounding the sensory dendrites. In the males of all three species, the pheromone-sensitive long sensilla trichodea exclusively contained PBP. the majority of the sensilla basiconica in both sexes in these species contained GOBP; these sensilla are known to respond to plant and other general odours. Some sensilla were not labelled by either antiserum; presumably, these held an odorantbinding protein of a different subfamily. Never were PBP and GOBP co-localized in the same sensillum. Two observations deserve special attention: (1) PBP was also found in a few sensilla in females, and (2) in B. mori, where the long sensilla trichodea have a different functional specificity in males (pheromone) and females (plant odours), the expression of the odorant-binding protein (males: PBP; females: GOBP) is similarly different. The distinct and complex distribution pattern of odorant-binding proteins supports the notion that these proteins participate in stimulus recognition.Dedicated to Professor Ya.A. Vinnikov on the occasion of his 85. birthdayThis work was partly supported by DFG grant ste 501/3-1.     

Olfaction in the Colorado potato beetle: Ultrastructure of antennal sensilla inLeptinotarsa sp

Sensillae on the antennae of the Colorado potato beetle,Leptinotarsa decemlineata are described using scanning (SEM) and transmission (TEM) electron microscopy and compared with SEM observations of antennal sensilla inL. haldemani andL. texana. In all the three species, 13 distinct sensillar types were identified with a higher density of sensilla in the more polyphagous species,L. decemlineata than in the moderately host specificL. haldemani and the highly host specificL. texana. Cuticular specializations and the predominance of olfactory sensilla are discussed in relation to host specificity in the three species.