Release of male courtship display in Periplaneta americana: evidence for female contact sex pheromone

Male American cockroaches (Periplaneta americana) are attracted to virgin females by volatile sex pheromones. After antennal contact with the female they turn through 180° and spread their wings in courtship display. A chemical contact stimulus releasing male courtship is demonstrated in the female cuticle. Experiments with standardized olfactory stimulation by volatile sex pheromones revealed that the contact stimulus is sex-specific and species-specific. It can be washed off the cuticle with non-polar solvents and was successfully transferred to glass dummies. However, it is not effective in the absence of volatile sex pheromones. Thus volatile sex pheromones are responsible for male attraction and sexual motivation, while mate recognition is accomplished through the contact pheromone.     

Sexual dimorphism of tarsal receptors and sensory equipment of the ovipositor in the European corn borer,Ostrinia nubilalis

Summary Sensilla on legs and ovipositor of the moth Ostrinia nubilalis were investigated by light and scanning electron microscopy. The ovipositor is composed of two papillae densely packed with medium length mechanoreceptor sensilla (MRb: 80–160 m, n=420–460). Long mechanoreceptor sensilla (MRa: 250–300 m, n=20–24) and contact chemoreceptors (CRa: 30–40 m, n=20–28) are evenly distributed at the periphery of these papillae. Legs support contact chemoreceptors (CRa), scattered among the scales. The pretarsus structure of each leg includes a single contact chemoreceptor (125 m) inserted dorsally. The fifth tarsomere bears a ventral area without scales on which contact chemoreceptors are disposed in two parallel rows (CRb: 40–60 m). A sexual dimorphism was found in the number and density of these sensilla (females: mean = 5.3, SD=1.0; males: mean = 3.3, SD=0.7), and in the size of the sensory field. The possible involvement of these sensory structures in oviposition site selection is discussed.     

Flagellar setae of the second antennae in decapod shrimps: sexual dimorphism and possible role in detection of contact sex pheromones

Summary

The antenna 2 (antennal) flagella of decapod shrimps are chemotactile, and their setae are proposed as sensilla involved in recognition of females by males via a contact sex pheromone on the surface of the female. Male recognition of females receptive to mating occurs in many caridean species upon contact of male antennal flagella with the surface of a newly molted parturial female. The hypothesis of sexual dimorphism in the number and kind of setae on the antennal flagella of four caridean and one penaeid shrimp species was tested with setal counts and observations on setal morphology. Unique male antennal setae (“male-specific sensilla”) were not identified in any of the species investigated. However, the abundance of antennal setae was significantly greater in males than in breeding females in the palaemonid carideans Palemonetes pugio and Macrobrachium ohione. In the hippolytid caridean Thor manningi and alpheid caridean Alpheus normanni, no sexual dimorphism in setal abundance was demonstrated. In the penaeoid Rimapenaeus similis, males had a higher abundance of antennal setae than the larger breeding females but so did juvenile females, similar in size to males. The sexual dimorphism in antennal sensilla in the palaemonid species and its absence in A. normanni might be related to their different mating systems, but no such association is suggested for T. manningi and R. similis. Setal morphology suggestive of chemoreceptive function (a terminal pore) was observed in all species.     

Tubular bodies in dendritic outer segments projecting to second embryonic cuticle from anlagen of contact chemoreceptors in Locusta migratoria L. (Orthoptera : Acrididae) and Periplaneta americana (L.) (Dictyoptera : Blattidae)

Bundles of dendritic outer segments project to the 2nd embryonic cuticle from anlagen of contact chemoreceptors in the antennae of 9-day embryos (80–85% stage) of Locusta migratoria L. (Orthopteroidea) and in the maxillary palps of 24-day embryos (80% stage) of Periplaneta americana (L.) (Blattodea). The dendrite sheaths fuse with irregular nodular thickenings in the 2nd embryonic cuticle in Locusta; in Periplaneta the sheaths are joined to it by filaments. In both species one of the dendritic outer segments shows a dilation within which a tubular body is contained. This tubular body is located in Locusta at the nodular thickening where the dendrite sheath is connected to the 2nd embryonic cuticle. In Periplaneta, the tubular body was observed near the junction between the dendrite sheath and the cuticle in most cases, but at some distance to the cuticle in others. The occurrence of a tubular body is discussed with reference to mechanoreceptive capacities of the sensory cells and to the phylogeny of the 2nd embryonic cuticle.     

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.     

Les sensilles de l'antenne chez quelques blattes et en particulier chez Blaberus craniifer (Burm.)

Résumé Les organes sensoriels du scape et du pédicelle sont essentiellement propriocepteurs. Le fouet est très riche en chimiorécepteurs (plus de 100000) mais chez Periplaneta et Blatta, on observe un net dimorphisme sexuel qui n'existe pas chez Blaberus. La relation entre un tel dimorphisme et la perception des phéromones sexuelles est discutée. L'ultrastructure des trois principaux types de sensilles est décrite: Ce sont des grandes soies gustatives et mécanoréceptrices, des petites soies olfactives à paroi mince et percée de pores et de tubules qui sont innervées par un ou deux neurones, des petites soies olfactives à paroi épaisse et striée qui contiennent toujours quatre neurones. L'étude électrophysiologique préliminaire confirme les résultats ci dessus en ce qui concerne le rôle des sensilles: La phéromone grégaire peut être perçue au niveau de certaines petites soies olfactives striées.

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The antennal sensilla of some cockroaches with special reference to Blaberus craniifer (Burm.)

Summary The antennae of cockroaches were studied with both scanning and transmission electron microscope. The sense organs of the scape and pedicel are essentially proprioceptors. The chemoreceptors are very numerous on the flagellum (more than 100000) but in the case of Periplaneta and Blatta, a significant sexuel dimorphism is noticed. Such a dimorphism does not exist in Blaberus and its relation with the perception of the sexual attractants is discussed. Three principal sense organs of Blaberus are described: big thick walled hairs with gustatory and mechanoreceptive function; thin walled olfactory hairs with pore-tubules innervated by one or two neurons; thick walled grooved hairs with four neurons and olfactory function. Special attention was given to the secretory aspect of the accessory cells in the olfactory sensilla. A preliminary electrophysiological investigation confirms the above mentioned functional interpretations. The gregarious pheromone seems to be perceived by some little thick walled grooved hairs.

     

Population genetic structure and intraspecific genetic distance of Periplaneta americana (Blattodea: Blattidae) based on mitochondrial and nuclear DNA markers

The American cockroach (Periplaneta americana) is a globally invasive pest that can cause significant economic loss and threaten human health. Although it is abundant and lives in close proximity to humans, few studies have investigated the genetic diversity of P. americana. Our study analyzed 1,053 P. americana and other Periplaneta species' samples from different locations in China and the United States. A traditional tree‐based method using 17 unique mitochondrial COI haplotypes of P. americana and 20 haplotypes of the other Periplaneta species accurately identified P. americana with a barcoding threshold of 5.1%. To identify the population genetic structure of P. americana, we investigated wingless gene and pooled them with obtained mtDNA data for a combined analysis. Although the genetic diversity of the USA group was relatively higher than the China group, the number of haplotypes and alleles of both groups was small. The analysis of molecular variance (AMOVA), intraspecific phylogeny, and haplotype networks indicated that P. americana had very little global genetic differentiation. The weak geographic genetic structure might reflect the human‐mediated dispersal of P. americana. Despite no apparent phylogeographic assignment of mtDNA and nuclear lineages was observed in both BI trees, the integrated COI sequence data identified four distinct P. americana haplotype groups, showing four ancient maternal lineages of P. americana in China and the United States.     

The structure of the antennae of the Florida Queen butterfly, Danaus gilippus berenice (Cramer)

Interest in the structure of the antennae of the Florida Queen butterfly arises from the finding that a pheromone is active in their courtship. Light and electron microscopic techniques were used to study the sensilla on the antennae and three types of sensilla with perforated walls were identified. The most common of these are short, thin-walled pegs which are distributed over most of the antennal surface. Long, curved, thin-walled pegs occur in patches on the inner medial antennal surface. Multiple coeloconic sensilla are present having up to 50 pegs in one sensillum. On the outer 28 flagellar subsegments there are two such sensilla per subsegment. In addition there are on the antennae long, thick-walled hairs which are mechanoreceptors and probably also contact chemoreceptors. Sunken pegs, the function of which is not known, occur on the antennae. Grooved sensilla were found with the electron microscope but could not be identified with the light microscope. There was no indication of sexual dimorphism in sensilla types or numbers on the antennae.     

Ultrastructure of antennal sense organs of small ermine moths,Yponomeuta spp. (Lepidoptera : Yponomeutidae)

The ultrastructure of antennal sensilla was investigated in males and females of 5 European Yponomeuta species. Three types of olfactory sensilla could be distinguished, i.e. sensilla trichodea, sensilla basiconica, and sensilla coeloconica. Sexual dimorphism in the number of cells innervating sensilla trichodea could be established in 3 species. Most of these sensilla in males of Y. vigintipunctatus, Y. rorellus, and Y. cagnagellus are associated with 3 sensory cells, whereas in females of these species these sensilla contain 2 sensory cells. In both males and females of Y. padeltus and Y. malinellus, the great majority of the sensilla trichodea is innervated by 3 cells. So-called sensilla chaetica type 1 show features typical of a combined mechano/contact-chemoreceptor. No structures indicating a sensory function were found in sensilla chaetica type 11. Sensilla styloconica contain 3 cells, the dendritic outer segment of one being lamellated; most likely, these cells have a thermo- and hygroreceptive function. The findings concerning the olfactory sensilla are discussed in the context of olfactory communication and reproductive isolation in European Yponomeuta.     

Antennal sensilla of male and female of the nut weevil,Curculio nucum Linnaeus, 1758 (Coleoptera: Curculionidae)

The nut weevil, Curculio nucum (Linnaeus, 1758), is the main pest in hazelnut orchards (Corylus avellana L.). Semiochemicals are interesting bio control tools that could be used to manipulate the pest behaviour and to control pest populations. The study of the sensorial equipment of the insect antennae provides information on the importance of olfaction in the adult life for host plants and mate findings as well as on the putative other senses. Before electrophysiological investigation, the knowledge of antennae equipment is also necessary. The aim of this study is to determine the types, number and location of sensilla on the antennae of male and female adult C. nucum in order to determine their implication in seeking a sexual partner and a host plant. The 12-segmented antenna comprises a scape, a 7-segmented funicle and a 4-segmented club. Out of the nine sensillum types listed, three are present on the scape and the funicle and seven types on the club which gathers 71–73% of the total of sensilla. Tactile aporous sensilla chaetica C1, gustatory uniporous sensilla chaetica C2, olfactory multiporous sensilla basiconica B1 and B2 are found on both the dorsal and ventral surfaces of the club in both sexes. Thermo-hygroreceptive dome-shaped sensilla D, olfactory multiporous sensilla basiconica B3 and olfactory multiporous fluted sensilla basiconica F are found exclusively on the ventral surface of the club, suggesting that these sensilla are utilized in host plant acceptance during antennal tapping. The sexual dimorphism concerns only the numbers of sensilla chaetica C1 and sensilla basiconica B2.     

Sucrose-binding lectin in regenerating cockroach (Periplaneta americana) legs: Its purification from adult hemolymph

Previously, we purified Periplaneta lectin from the hemolymph of adult Periplaneta americana (American cockroach) (Kubo and Natori Eur. J. Biochem. 168, 75–82, 1987). Immunoblotting analysis using antibody against Periplaneta lectin showed that the cockroach hemolymph contains another lectin that cross reacted immunologically with Periplaneta lectin. We have purified this new lectin (regenectin) to homogeneity. Affinity purified antibody against regenectin cross reacted with Periplaneta lectin. Thus, Periplaneta lectin and this new lectin were found to be different lectins sharing common antigenicity. After leg amputation, this new lectin was found to appear transiently at a specific stage of regeneration as revealed by immunoblotting, suggesting that it plays a role in the regeneration process.     

Sex-specific non-pheromonal taste receptors in Drosophila

Taste receptorshave recently been reported in Drosophila [1] and [2], but little is known of the relation between receptor and response. Morphological studies of the distribution of chemosensory sensilla indicate that the fruit fly has two major sites of gustation: the proboscis and the legs [3]. The taste sensilla on both these sites are similar in structure and each sensillum generally houses four gustatory neurons [4]. Early anatomical observations have demonstrated a sexual dimorphism in the number of tarsal sensilla [5] and in their central projections [6]. We measured the electrophysiological responses of the prothoracic taste sensilla to non-pheromonal substances—salts, sugars and water—and found a clear sexual dimorphism. From the response profile of individual sensilla, we were able to distinguish three types of tarsal sensilla in females as against only two types in males. The female-specific type, which responded specifically to sugar, was absent in males except when male gustatory neurons were genetically feminised. The fact that tarsal gustatory hairs exhibit a sexual dimorphism that affects the perception of non-pheromonal compounds suggests that sexual identity is more complex than has previously been thought [7] and [8].     

The antennae and antennal sensilla of the giant looper,Boarmia (Ascotis) selenaria SchiffermÜller (Lepidoptera,Geometridae)*

The general morphology of the antennae and the antennal sensilla of both sexes of Boarmia (Ascotis) selenaria SchiffermÜller are described. The following types of sensilla were found: Bohm bristles and sensilla chaetica, trichodea, coeloconica, auricillica and styloconica. Antennal sexual dimorphism is marked, but mainly confined to the presence of long sensilla trichodea in males and the absence in females. The sensilla trichodea are the most abundant sensillum type.     

External morphology of antennae and their sense organs in the roach Gromphadorhina brunneri (Blattoidea: Dictyoptera)

The antennae and their sense organs in nymphs and adult roaches of Gromphadorhina brunneri, were investigated and described. The number of segments and sensillae of the nymphal antennae depend on the developmental stage. Sexual dimorphism is pronounced. Males have longer antennae than females as well as an abundance of especially long sensory hairs (long wavy hairs), which are probably responsible for the perception of female sex pheromones. They also have more thin-walled sensory hairs, for instance, sensilla trichodea. On a morphological basis the sensillae of Gromphadorhina brunneri, were named and classified. Long wavy hairs and large sensory hairs appear to be present also in a related species, G. portentosa, but are lacking in others. Their distribution on the antennae varies greatly from that in G. portentosa but their structure varies only slightly. These two types of sense organs are considered to be specialized forms of sensilla chaetica. They are contact chemoreceptors, as are two other types of sensilla chaetica. Furthermore, thin-walled chemoreceptors are present, such as sensilla trichodea, sensilla basiconica, sensilla coeloconica and a typical mechanoreceptor, the sensillum campaniformium.     

Environmental effects on numbers of peripheral chemoreceptors on the antennae of a grasshopper

Rearing conditions affected the numbers of chemoreceptor sensillaon the antennae of the grasshopper, Schistocerca americana.The numbers of olfactory sensilla were increased when insectswere reared in less complex odor environments (in isolationv. in a crowd, on a single food plant v. a mixture of plants,in an ‘odor-free’ environment v. a highly odorousenvironment). Numbers of contact chemoreceptors were not affectedby crowding, but were lower in insects from the single plantand ‘odor-free’ environments. The differences variedalong the antenna with insects from the less complex environmentshaving relatively more sensilla on the distal annuli and similarnumbers, or even fewer, on the proximal annuli. Numbers of sensillain the pre-oral cavity were not affected by rearing conditions.     

Sexually dimorphic antennal structures of New Zealand Cave Wētā (Orthoptera: Rhaphidophoridae)

Insect antennae carry diverse sensory organs for detecting information about their environments and facilitating interaction among individuals, particularly for reproductive behaviour. As such they are predicted to be sexually dimorphic, especially in species which do not use visual or auditory mate-location. The cave wētā are such animals, suggesting that they rely heavily on chemical and tactile information to interact. Sensilla morphology and sexual dimorphism was investigated in two species; Pachyrhamma acanthocera and P. waitomoensis, using scanning electron microscopy. Fourteen types of sensilla basiconica, campaniformia, chaetica, coeloconica, trichodea and Böhm’s sensilla were identified. Among these, sexual dimorphism in the length of four subtypes was detected, and surprisingly, P. waitomoensis males completely lacked all but two of the porous sensilla types. In addition, three types of unique horn-like and hooked antennal protrusions in male P. acanthocera are described, along with similar, but less diverse structures on the antennae of male P. waitomoensis.     

Fine structure of tarsal sensory organs in the whip spider Admetus pumilio (Amblypygi, Arachnida).

The sensory organs on the tarsi of the antenniform first legs of the whip spider Admetus pumilio C. L. Koch (Amblypygi, Arachnida) were examined with the scanning and transmission electron microscope. At least four different types of hair sensilla were found: (1) thick-walled bristles, which have the characteristics of contact chemoreceptors (several chemoreceptive dendrites in the lumen plus two mechanoreceptors at the base); (2) short club sensilla, innervated by 4-6 neurons which terminate in a pore on the tip; they are possibly humidity receptors; (3) porous sensilla, which are either innervated by 20-25 neurons and have typical pore tubules, or they have 40-45 neurons but no pore tubules; both types are considered to be olfactory; (4) rod sensilla occur in clusters near segmental borders; they are innervated by only one large dendrite which branches inside the lumen. Other tarsal receptors are the claws, which correspond to contact chemoreceptors, and the pit organ which resembles the tarsal organ of spiders. Compared to other arthropod sensilla, the contact chemoreceptors are very similar to those of spiders, while the porous sensilla correspond structurally to olfactory receptors in insects; the club and rod sensilla seem to be typical for amblypygids.     

Lectin staining of the dendrite sheath in contact chemosensitive sensilla of Periplaneta americana

The two sensillum lymph cavities of contact chemosensitive sensilla of insects are separated by the dendrite sheath. The composition of the dendrite sheath in the contact chemosensitive sensilla of the ventral sensory field on the maxillary palps of Periplaneta americana L. has been studied using lectin cytochemistry. The dendrite sheaths can be isolated from the surrounding tissue and the autofluorescent cuticle by incubation of cryostat sections in phosphate-buffered saline containing 1% Triton X-100. Various lectins bind specifically to the dendrite sheath indicating the presence of sugar residues. Enzymatic digestion with hyaluronidase or trypsin prior to lectin cytochemistry decreases staining intensity, whereas treatment with neuraminidase and chitinase causes no visible change. The material of the dendrite sheath can thus be assumed to contain acid mucopolysaccharides that probably are similar to those of the receptor lymph cavities. The negatively charged, hydrophilic acid mucopolysaccharides may make the dendrite sheath permeable for small cations. These findings are discussed in the context of the functional significance of the dendrite sheath in contact chemosensory sensilla of insects.     

Changes in distribution and abundance of antennal sensilla during growth of Locusta migratoria L. (Orthoptera : Acrididae)

The antenna of Locusta migratoria (Orthoptera : Acrididae) increases in length by the production of new annuli proximally and by elongation of existing annuli. The most distal annuli are fully differentiated at the time of hatching and no new olfactory sensilla are added to them at subsequent molts. More proximally, the differentiation of trichoid contact chemoreceptors precedes the development of olfactory sensilla. Sensillum differentiation proceeds from distal to proximal along the antenna and more distal annuli attain full development at each molt. The biggest increase in numbers of olfactory sensilla occurs at the final molt. On any one annulus, most olfactory sensilla are restricted to sensory fields on the anterior and posterior faces. Their spacing within these fields is consistent with the existence of separate but interacting chaetogens regulating the differentiation of basiconic and coeloconic sensilla.     

Response of <Emphasis Type="Italic">Trichogramma chilonis</Emphasis> to infochemicals: an SEM and electrophysiological investigation

The external morphology and peripheral olfactory responses of antennal chemoreceptors of Trichogramma chilonis Ishii (Hymenoptera:Trichogrammatidae) to host related stimuli were examined by SEM and EAG technique respectively. The antennae of male and female T. chilonis reveal strong sexual dimorphism in structure as well as in types, density and distribution of individual sensillar types. The female antennae are characterized by the presence of a single uniporous sensilla on the apical tip of the dome and numerous multiporous pitted sensilla. Sensilla common to both males and females include the multiporous plate sensilla, the peg sensilla and trichoid sensilla. Differences between males and females occur with respect to their EAG profiles to 22 chemicals commonly found in plants viz., hexanal, -pinene, citronellal, terpinene, citronellol, phytol, caryophyllene, R-(+)-limonene, linalool, carvacrol, cis-3-hexen-1-ol, ethanol, -pinene, trans-2-hexanal, -phellandrene, amyl acetate, cineole, quinolene, S-(–)limonene, -myrcene, carvone, terpineol; hexane leaf extracts of three host plants of Helicoverpa armigera (cotton, chickpea and pigeon pea) and hexane scale extracts of adult males and females of H. armigera. Individually, the largest peak amplitudes in females were obtained with citronellal, phytol, caryophyllene, R-(+)-limonene, linalool, carvacrol and citronellol while in males citronellol, caryophyllene, linalool, R-(+)-limonene and amyl acetate caused the highest response. Among the different chemical classes, the largest peak amplitudes in females were obtained with oxygenated monoterpenes, while in males, fatty acid derivatives caused the highest response. In both males and females, the hydrocarbon monoterpenes caused the least response. Thus, the wide spectrum response of Trichogramma chilonis, to a wide group of chemicals belonging to different chemical classes, though varying, nevertheless suggests that using plant information enhances its parasitization efficiency.