Adult Male Antennae Research Articles

Antennal Sensitivity of Spotted Lanternflies, Lycorma delicatula: Differential Electrophysiological Responses of Males and Females to Compounds Derived from Host Plants and Conspecifics.

In herbivorous insects, antennae play a crucial role in chemical communication and orientation when locating host plants and mates. To evaluate antennal sensitivity in response to odor stimuli, electroantennography (EAG) has been a practical technique. In the current study of the invasive spotted lanternfly (SLF), Lycorma delicatula (Hemiptera: Fulgoridae), we evaluated and compared their antennal sensitivity to a series of volatile chemicals collected from their bodies, honeydew, and host plants. To do this, we exposed the antennae of SLF fourth-instar and adult males and females to individual chemicals at a fixed dose of 50 ng. Further, a series of dose-response tests were carried out within a range of 0.5 to 100 ng. Although the amplitude of antennal responses varied among stages and sexes, adult males generated the strongest antennal responses in both experiments. In dose-response experiments, increased doses of chemicals up to 50 ng revealed the saturation points except in adult females which required a higher dose (100 ng) to reveal the saturation point. Although EAG does not provide any information on behavioral responses, our results are consistent with the olfactory bioassays in previous publications in which adult males, not females, were attracted to natural volatiles of their conspecifics. EAG indicated a higher sensitivity of adult male antennae to odor stimuli, particularly conspecific volatiles, than female antennae and highlighted sexual differences in the perception of chemical cues in SLF.

The structure and morphogenic changes of antennae of Spilococcus alhagi (Hall) (Hemiptera: Coccoidea: Pseudococcidae) in different instars

To better understand the antennal morphology and sensilla distribution of Spilococcus alhagi (Hall) in different instars, the antennae are examined using light microscopy and scanning electron microscopy. The results show that the antennae of S. alhagi could be differentiated in different instars based on their stucture, number of flagellomere, and sensillar distribution. Functional antennae of S. alhagi appear in first instar nymphs, adult females, and adult males as many scale insects. The antennae of first-instar nymphs comprise six segments, and each antenna has 62–71 sensilla of eight types, including sensilla basiconica (SB), one type of sensilla trichoidea (ST1), two types of sensilla companiform (SCa, sSCa), two types of sensilla coeloconica (SCo1, SCo2) and two types of sensilla chaetica (SCh1, SCh2). The antennae of adult females have eight segments, but each antenna has 164–188 sensilla of the same eight types as the first instar nymphs. The antennae of adult males are more developed, and each antenna contains ten segments and 670–725 sensilla of eleven types, including knobbed seta sensilla (Ks), intersegmental sensilla (Ins), two types of sensilla trichoidea (ST1, ST2), two types of sensilla companiform (SCa, sSCa), two types of sensilla coeloconica (SCo1, SCo2) and three types of sensilla chaetica (SCh1, SCh2, bSCh). Of these, four types of sensilla, viz., Ks, ST2, Ins, bSCh are only observed on the antennae of adult males. The prepupa-like and pupa-like male nymphs both possess simplified antennae, and exhibit four types of sensilla (ST1, SCa, SCh1, SCh2). Based on these results, the morphogenic changes and potential functions of antennae of S. alhagi in different instars are discussed.

Chemosensory protein 4 is required for Bradysia odoriphaga to be olfactory attracted to sulfur compounds released from Chinese chives.

Bradysia odoriphaga (Diptera: Sciaridae) is a serious pest of Chinese chives cultivated in China. Chemosensory proteins (CSPs) are important components of insect olfactory systems that capture and bind environmental semiochemicals which are then transported to olfactory receptors. Despite their importance, the mechanism of olfaction and related behavioral processes in B. odoriphaga have not been characterized. Here, we found that BodoCSP4 has an important olfactory function. RT-qPCR indicated that BodoCSP4 expression was highest in the heads (antennae removed) of adult males, followed by the antennae of adult males. Competitive binding assays with 33 ligands indicated that BodoCSP4 binds well with methyl allyl disulfide, diallyl disulfide, and n-heptadecane; the corresponding dissolution constants (Ki) were as high as 5.71, 5.71, and 6.85 μM, respectively. 3D-structural and molecular docking indicated that BodoCSP4 has five α-helices and surrounds the ligand with certain hydrophobic residues including Leu60, Leu63, Leu64, Ala67, Val28, Ile30, Ile33, Leu34, and Val86, suggesting these residues help BodoCSP4 bind to ligands. Silencing of BodoCSP4 significantly decreased the attraction of B. odoriphaga males to diallyl disulfide and n-heptadecane but not to methyl allyl disulfide in Y-tube olfaction assays. These results increase our understanding of how BodoCSP4 contributes to host and female localization by B. odoriphaga males.

Silencing the odorant coreceptor (Orco) disrupts sex pheromonal communication and feeding responses in Blattella germanica: toward an alternative target for controlling insect-transmitted human diseases.

The German cockroach, Blattella germanica, is one of the most severe pests of urban and rural areas. High-throughput genetic screening approaches indicate that the olfactory system of this pest is extremely powerful because it has an extensive array of olfactory receptor genes compared with many other insect species. Several of these genes have been identified previously, but their functions have not yet been characterized. This study describes the sequence of five transcriptomes of B. germanica adult male antennae, female antennae, maxillary palps, legs, and fifth-instar nymph antennae to investigate expression patterns of odorant receptors (ORs). Approximately 90% of ORs were found to be the most highly expressed genes in adult or nymph antennae. Additionally, every OR requires an odorant co-receptor (Orco) to become fully functional, and this was selected and successfully inhibited by injection of the corresponding double-stranded (ds)RNA targeting the Orco. A strong RNA interference (RNAi) effect was observed in which > 75% of Orco messenger RNA (mRNA) was clearly suppressed after 72 h of treatment. Olfactory behavioral assays showed that Orco-impaired B. germanica respond more slowly and show less attraction to one volatile sex pheromone and food resources compared with a control group. The results show that Orco plays a pivotal role in both sex pheromone and food-seeking olfactory processes, and provide an alternative genetic technique for controlling this urban pest species by olfactory disruption. © 2020 Society of Chemical Industry.

Olfactory Proteins in Timema Stick Insects

Most animals use olfaction to obtain important information from the environment, including the presence of food or mates. Insects detect odorants through receptors that are expressed in the sensory neurons of the olfactory sensilla, which cover the surface of the antennae. The olfactory capacities of an insect thus depend largely on the repertoire of the odorant receptors. Here, we study the repertoire of olfactory proteins in the stick insect Timema cristinae. We first generate transcriptomes from the antennae of adult males and females and identify, via homology searches, putative olfactory proteins of three different families: odorant binding proteins, odorant receptors, and chemosensory proteins (CSPs). We then attempt to categorize olfactory proteins likely involved in sexual communication by comparing gene expression between adults and juveniles, as well as between males and females. Notably, the olfactory proteins involved in the perception of food or abiotic environmental components, should be expressed in both adults and juveniles. By contrast, the olfactory proteins involved in sexual communication, such as the detection of sex pheromones, should be expressed in adults and often comprise different repertoires in males and females. Finally, we also tested whether olfactory proteins in general and the subset, with putative roles in sexual communication in particular, are under relaxed selection in the asexual species T. monikensis, a close relative of T. cristinae. We found that olfactory proteins are typically differentially expressed between juveniles and adults, but there is little overlap of differential expression between developmental stages and the level of sex bias in adults. Furthermore, while we find evidence that olfactory proteins are indeed under relaxed selection in the asexual species, there is no evidence that this is necessarily the case for olfactory genes with a putative role in sexual communication. Nevertheless, the list of olfactory genes generated in our study provides a useful tool for future studies on olfaction in Timema and other stick insects.

Antennal sensilla of Didesmococcus koreanus Borchsenius (Hemiptera: Coccoidea: Coccidae) in different instars

To better understand the morphology, structure and function of antennae and antennal sensilla of Didesmococcus koreanus Borchsenius, family Coccidae (Hemiptera: Coccoidea), in the first-instar nymph, adult female and adult male stages, we investigated these stages using light microscopy and scanning and transmission electron microscopy. The results show that the antennae of first-instar nymphs and adult females are similar in morphology and sensilla distribution. The antennae of first-instar nymphs comprise six segments, and each antenna has 24 sensilla of six types: one campaniform sensillum (Ca), eleven smooth aporous trichoid sensilla (St), two basiconic sensilla (Ba), five multiporous pegs (Mp), two smooth multiporous pegs (sMp) and three chaetica sensilla (Ch). The antennae of adult females have six or seven segments, and each antenna has 25 sensilla of the same six types as the first nymphs, but with twelve sensillum St. The antennae of adult males are more developed, and each antenna contains 155–213 sensilla of eight types: 16–34 St, 1 Ca, 2–5 Ba, 1 Johnston’s organ (Jo), 3 knob sensilla (Ks), 5 Mp, 2 sMp and 120–181 multiporous sensilla trichodea (Mt). Based on our analysis, we discuss the possible functions of the antennae and antennal sensilla of D. koreanus in different instars.

Larval sensilla of the moth Heliothis virescens respond to sex pheromone components.

Female-released sex pheromones orchestrate the mating behaviour of moths. Recent studies have shown that sex pheromones not only attract adult males but also caterpillars. Single sensillum recordings revealed that larval antennal sensilla of the moth Heliothis virescens respond to specific sex pheromone components. In search for the molecular basis of pheromone detection in larvae, we found that olfactory sensilla on the larval antennae are equipped with the same molecular elements that mediate sex pheromone detection in adult male moths, including the Heliothis virescens receptors 6 (HR6) and HR13, as well as sensory neurone membrane protein 1 (SNMP1). Thirty-eight olfactory sensory neurones were identified in three large sensilla basiconica; six of these are considered as candidate pheromone responsive cells based on the expression of SNMP1. The pheromone receptor HR6 was found to be expressed in two cells and the receptor HR13 in three cells. These putative pheromone responsive neurones were accompanied by cells expressing pheromone-binding protein 1 (PBP1) and PBP2. The results indicate that the responsiveness of larval sensilla to female-emitted sex pheromones is based on the same molecular machinery as in the antennae of adult males.

The structure and morphogenic changes of antennae of Matsucoccus matsumurae (Hemiptera: Coccoidea: Matsucoccidae) in different instars

To better understand the functioning and morphogenic changes of the antennae of Matsucoccus matsumurae (Kuwana) in different instars, the antennae are examined using light microscopy, scanning and transmission electron microscopy. The results show that the antennae of M. matsumurae display three different styles in morphology and sensillar distribution in different instars. The antennae of first instar nymphs are relatively simple, including one campaniform sensillum (Ca), four smooth aporous trichoid sensilla (SAt), two intersegmental sensilla (Ins), two coeloconic sensilla (Co), three multiporous pegs (Mp) and four uniporous pegs (Up). The antennae of adult females and third instar male nymphs both possess similar antennae, and exhibit seven types of sensilla. Adult female antennae have in total 82–108 sensilla, including 9–16 Böhm's bristle (Bb), 3–7 Ca, 50–75 SAt, 0–3 Ins, 3–10 Co, 8 Mp and 5 Up, whereas third instar male nymph antennae possess approximately 62–79 sensilla. Adult male antennae are the most developed, possessing 259–312 sensilla, including 7–15 Bb, 2–5 Ca, 7–11 grooved aporous trichoid sensilla, 4–9 SAt, 0–3 Ins, 2–7 Co, 23–29 knobbed seta sensilla, 179–230 multiporous trichoid sensilla and 8 Mp. Based on these results, the main functions and morphogenic changes of antennae M. matsumurae in different instars are discussed.

Endogenous fatty acids in olfactory hairs influence pheromone binding protein structure and function in Lymantria dispar

The gypsy moth utilizes a pheromone, (7R,8S)-2-methyl-7,8-epoxyoctadecane, for mate location. The pheromone is detected by sensory hairs (sensilla) on the antennae of adult males. Sensilla contain the dendrites of olfactory neurons bathed in lymph, which contains pheromone binding proteins (PBPs). We have extracted and identified free fatty acids from lymph of sensory hairs, and we demonstrate that these function as endogenous ligands for gypsy moth PBP1 and PBP2. Homology modeling of both PBPs, and docking of fatty acids reveal multiple binding sites: one internal, the others external. Pheromone binding assays suggest that these fatty acids increase PBP-pheromone binding affinity. We show that fatty acid binding causes an increase in α-helix content in the N-terminal domain, but not in the C-terminal peptide of both proteins. The C-terminal peptide was shown to form a α-helix in a hydrophobic, homogeneous environment, but not in the presence of fatty acid micelles. Through partition assays we show that the fatty acids prevent adsorption of the pheromone on hydrophobic surfaces and facilitate pheromone partition into an aqueous phase. We propose that lymph is an emulsion of fatty acids and PBP that influence each other and thereby control the partition equilibria of hydrophobic odorants.

(R)-Desmolactone is a sex pheromone or sex attractant for the endangered valley elderberry longhorn beetle Desmocerus californicus dimorphus and several congeners (Cerambycidae: Lepturinae).

We report here that (4R,9Z)-hexadec-9-en-4-olide [(R)-desmolactone] is a sex attractant or sex pheromone for multiple species and subspecies in the cerambycid genus Desmocerus. This compound was previously identified as a female-produced sex attractant pheromone of Desmocerus californicus californicus. Headspace volatiles from female Desmocerus aureipennis aureipennis contained (R)-desmolactone, and the antennae of adult males of two species responded strongly to synthetic (R)-desmolactone in coupled gas chromatography-electroantennogram analyses. In field bioassays in California, Oregon, and British Columbia, traps baited with synthetic (R)-desmolactone captured males of several Desmocerus species and subspecies. Only male beetles were captured, indicating that this compound acts as a sex-specific attractant, rather than as a signal for aggregation. In targeted field bioassays, males of the US federally threatened subspecies Desmocerus californicus dimorphus responded to the synthetic attractant in a dose dependent manner. Our results represent the first example of a “generic” sex pheromone used by multiple species in the subfamily Lepturinae, and demonstrate that pheromone-baited traps may be a sensitive and efficient method of monitoring the threatened species Desmocerus californicus dimorphus, commonly known as the valley elderberry longhorn beetle.

(R)-Desmolactone, A Female-produced Sex Pheromone Component of the Cerambycid Beetle Desmocerus californicus californicus (subfamily Lepturinae)

We report the identification, synthesis, and field bioassays of a female-produced sex attractant pheromone for the cerambycid beetle Desmocerus californicus californicus Horn. Headspace volatiles from females contained a sex-specific compound, (R)-desmolactone [(4R,9Z)-hexadec-9-en-4-olide], which elicited strong responses from the antennae of adult males in coupled gas chromatography-electroantennogram analyses. Short syntheses of both enantiomers were developed from commercial chiral synthons. In field bioassays, significant numbers of males were collected in traps baited with (R)-desmolactone, whereas the (S)-enantiomer attracted no males. The racemate was less attractive than the pure (R)-enantiomer, indicating some degree of antagonism by the unnatural enantiomer. This compound is the first example of a new structural class of cerambycid pheromones, and is the second pheromone identified for a species in the subfamily Lepturinae.

Cis-Vaccenyl Acetate, A Female-Produced Sex Pheromone Component of Ortholeptura valida, A Longhorned Beetle in the Subfamily Lepturinae

We report the identification, synthesis, and field bioassays of a female-produced sex attractant pheromone component of the cerambycid beetle Ortholeptura valida (LeConte). Headspace volatiles from females contained a female-specific compound, (Z)-11-octadecen-1-yl acetate, which elicited a strong response from antennae of adult males in coupled gas chromatography-electroantennogram analyses. In field bioassays, significant numbers of males were collected by traps baited with this compound. The pheromone represents a new structural class of cerambycid pheromones, and is the first pheromone identified for a cerambycid species in the subfamily Lepturinae.

Expressed sequence tags and proteomics of antennae from the tortricid moth, Epiphyas postvittana

Genomic and proteomic analyses of the antennae of the light brown apple moth, Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae) were undertaken to identify genes and proteins potentially involved in odorant and pheromone binding and turnover. An EST approach yielded 5739 sequences, comprising 808 contigs and 1545 singletons. InterPro and Blast analyses revealed members of families implicated in odorant and pheromone binding (PBPs, GOBPs, ABPXs and CSPs) and turnover (CXEs, GSTs, CYPs). Of the three pheromone binding proteins (PBPs) identified, two were more highly expressed at the RNA and protein levels in adult male antennae (EpPBP1, EpPBP3), while a third was more highly expressed in female antennae (EpPBP2). To identify proteins involved in the detection of sex-specific signals, differential 2D gel electrophoresis (pH 5-8) followed by mass spectrometry was conducted on antennal proteins from males versus females. Identified male-biased proteins included a pheromone binding protein, a porin, a short chain dehydrogenase/reductase, and a member of the takeout family.

Expression and localization of three G protein ? subunits, Go, Gq, and Gs, in adult antennae of the silkmoth (Bombyx mori)

In insect olfactory receptor neurons, rapid and transient increases in inositol triphosphate (IP3) and Ca2+ are detected upon stimulation with pheromone or nonpheromonal odorants. This suggests that heterotrimeric guanine nucleotide binding proteins (G proteins) may transduce some odorant responses in insects. We obtained cDNA clones encoding three classes of G protein alpha subunits, Bm Go, Bm Gq, and Bm Gs, from the antennae of the adult male silkmoth (Bombyx mori). RT-PCR experiments showed that the mRNA of these G protein alpha subunits was also present in the various tissues of adult and larval insects. We used immunocytochemistry to localize these G protein alpha subunits in adult male and female antennae. In the adult male antennae, anti-Go antiserum stained the nerve bundles. In contrast, anti-Gq and anti-Gs antisera stained the inner and outer dendritic segments of the putative olfactory receptor neuron. The localizations of Bm Go, Bm Gq, and Bm Gs in the female antennae were the same as in the male antennae. The localizations of Bm Gq and Bm Gs suggest that each subunit mediates a subset of the odorant response.

Intergeneric distribution and immunolocalization of a putative odorant-binding protein in true bugs (Hemiptera, Heteroptera).

Lygus antennal protein (LAP) is an olfactory-related protein of the tarnished plant bug Lygus lineolaris (Hemiptera, Heteroptera: Miridae), a hemimetabolous insect. In previous work, a polyclonal antiserum was generated against the N-terminal sequence of LAP; LAP immunoreactivity was strongest in antennae of adult males, but was also present in antennae of adult females and of nymphs. In the current study, LAP immunoreactivity was examined to determine the species specificity and the tissue and cellular localization of LAP expression. Western blot analysis indicated that LAP immunoreactivity was present in the antennae of the male congeners L. lineolaris and L. hesperous, but was not detectable in male antennae of the more distant relatives Podisus maculiventris or Nezara viridula (Hemiptera, Heteroptera: Pentatomidae). Western blot analysis further confirmed that LAP expression was restricted to antennal tissue. Histological analyses showed that LAP expression within the antennae was specifically associated with chemosensory sensilla on the antenna. Within the sensilla, LAP immunoreactivity was distributed throughout the extracellular lumen and was concentrated in dense granules within the cytoplasm of sensillar support cells. LAP immunoreactivity was restricted to a subset of antennal chemosensory sensilla, specifically the multiporous olfactory sensilla. These findings suggest that LAP has an important olfactory function in Lygus sp., possibly related to that of odorant-binding proteins (OBP) found in other insect orders. If so, LAP would be the first OBP-like protein characterized outside the Endopterygota.

Phosphorylation of cockroach antennal polypeptides: effects of second messengers and pheromonal blend

In insect antennal extracts, Schleicher et al.1 showed that protein kinase C (PKC) inhibitors abolish the transience of pheromone-induced rapid inositol trisphosphate responses, which suggests that pheromonal signals act on phosphorylation of specific proteins. To confirm this hypothesis, we studied the effects of second messengers and a pheromonal blend on phosphorylation of antennal proteins in the cockroachPeriplaneta americana. Proteins from adult male antennae were phosphorylated in vitro in the presence of [γ32P] triphosphate, then separated by SDS-polyacrylamide gel electrophoresis. Numerous phosphopolypeptides were visualized. The presence of Ca++/calmodulin in the incubation medium resulted in increased phosphorylation of polypeptides with molecular weights of 38, 48, 51, 54 and 58 kDa. Stimulation of PKC by addition of Ca++ phosphatidylserine (PS)/phorbol myristate acetate (PMA) resulted in the appearance of three phosphopolypeptides of 36, 70 and 120 kDa. In the presence of cyclic adenosine monophosphate, two new major polypeptides of 46 and 42 kDa appeared; the latter polypeptide also appeared in the presence of cyclic guanosine monophosphate. Comparison with polypeptide composition of tissue from the cerci, leg, brain and fat body showed that the 36 and 48 kDa polypeptides were specific to antennae, whereas the 120 kDa polypeptide was also present in the adult brain. When antennae are subjected to pheromonal stimulation for 16 seconds prior to homogenization, in vitro phosphorylation of the 120, 70, 64 and 38 kDa polypeptides was inhibited, whereas phosphorylation of the 58, 54, 51 and 48 kDa polypeptides was strongly stimulated. It is noteworthy that a 107 kDa polypeptide was observed only after pheromonal stimulation by Ca++/PS/PMA. Our findings suggest that Ca++-and PKC-dependent protein phosphorylation systems play an important role in the transduction of pheromonal signals in antennae of male cockroachP. americana. We speculate that specific phosphoproteins may modulate sensitivity and signal amplification during the olfactory transduction process.

Sex pheromone of caddisflyHesperophylax occidentalis (Banks) (Trichoptera: Limnephilidae)

The main component of the sex pheromone of the caddisflyHesperophylax occidentalis (Banks) (Trichoptera: Limnephilidae) was identified as 6-methylnonan-3-one (enantiomeric composition has not yet been determined). Extracts of adult females elicited strong electroantennogram (EAG) responses from adult male antennae, but elicited significantly smaller EAG responses from adult female antennae. Extracts of adult males did not elicit appreciable EAG responses from antennae of either sex. Racemic 6-methylnonan-3-one was synthesized and elicited EAG responses from male antennae as strong as those obtained with female extracts. In field tests with baited sticky traps near lakes and streams, traps baited with synthetic racemic 6-methylnonan-3-one caught significantly more males than control traps. Female adults contained approximately 1 µg of 6-methylnonan-3-one per individual. Related ketones and alcohols of other chain lengths were also tentatively identified, being present in tiny amounts in female extracts. Extraction of different body parts showed that 6-methylnonan-3-one occurs only in a region near the intersegmental membrane between the fourth and fifth abdominal sternites of the female (no discrete glands were observed). Extracts of males did not contain 6-methylnonan-3-one, nor did pupae of either sex.

Olfactory receptor responses of the nymphal American cockroach to sex pheromones and their mimics

1. 1. EAG responses to highly purified sex pheromones (periplanone-A and -B), sex pheromone mimics [germacrene-D, (+)-verbanyl acetate and (+)- trans-verbenyl acetate] and general odor (camphor) were recorded from both sexes of adult and three nymphal stages (7, 10 and 11th instars) of the American cockroach. 2. 2. The M/F ratios were evaluated for each stage by stimulation with the above chemicals. 3. 3. The ratio values indicated undeveloped sex pheromone receptor on the antennae of 7th male instar and females of all the stages. On the other hand, precursory development of the receptor was expected on the antennae of males of the old-aged nymphal instars and full development on the adult male antennae.

Sensilla of the antennae and the labial and maxillary palps of Blattella germanica (L.) (Dictyoptera: Blattellidae): Their classification and distribution.

The various sensilla on the antennae and on the labial and maxillary palps of Blattella germanica (L.) were studied. Thick-walled chemoreceptors with fluted shafts and articulated bases are located on the antennae and on the labial and maxillary palps. Thin-walled chemoreceptors, without fluted shafts or articulated bases, are restricted to the flagellar segments of the antennae and to the distal segments of the palps. Antennae of adult males have more thin-walled chemoreceptors than do those of females. Hair-plate sensilla are found at the scape-head and scape-pedicel joints, and at the joints of segments on the palps. Campaniform sensilla are concentrated as a ring around the distal margin of the pedicel, and are also scattered singly on the scape, pedicel, and flagellar segments of the antennae, and on the first segment of the maxillary palps. Occasionally, a few sensilla coeloconica and cold receptor sensilla are found on the antennal flagellum.

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.