Dimorphic antennal systems in gynandromorphic honey bees,Apis Mellifera l. (Hymenoptera: Apidae)

Gynandromorphic honey bees, Apis mellifera (Hymenoptera: Apidae), were examined to determine characteristic morphological and anatomical features of the antennal system. The antennae of gynandromorphic individuals are predominantly worker or drone-like. Hybrid antennae, composed of female and male tissues, occur only rarely (7 out of 188 examined antennae). Depending on the mosaic pattern of the head, both antennae can be drone-like or worker-like, or one can be drone-like and the other worker-like. Examination of the antennal lobes of six characteristic specimens revealed that antennal lobes, which are innervated by drone-like antennae, always have drone-specific enlarged tracts and macroglomerular complexes, whereas antennal lobes innervated by worker-like antennae always are composed of normally sized glomeruli. Thus, there is a strict correlation between the sexual morphology of the antennae and the sexual organization of the antennal lobe neuropil. In one antennal lobe, innervated by a hybrid antenna, we found a hypertrophied glomerulus, certainly homologous to one of the macroglomerular complexes in drone-like antennal lobes.jy 1999 Elsevier Science Ltd. All rights reserved.     

Electroantennogram responses to the sex pheromone and other odours in the American cockroach

Electroantennogram responses of adult American cockroaches were recorded for the sex pheromone and 30 kinds of monoterpenoids. Strong responses to the sex pheromone were obtained from the antennae of male adults, but not from females. The compounds possessing a hydroxyl or carbonyl group produced significant EAG responses in the antennae. Bornyl acetate, however, was an exception and gave strong responses in both male and female antennae. There was basically no difference between the antennal responsiveness to monoterpenoids tested in both sexes.     

Unusual pheromone receptor neuron responses in heliothine moth antennae derived from inter-species imaginal disc transplantation

Single-cell electrophysiological recordings were obtained from olfactory receptor neurons housed in sensilla trichodea along the adult antennae arising from transplantation of the antennal imaginal discs between larval male Helicoverpa zea and Heliothis virescens. The olfactory receptor neurons from the majority of type C sensilla sampled on transplanted antennae displayed response characteristics consistent with those of the species that donated the antennae. However, some of the sensilla type C sampled in either transplant type contained olfactory receptor neurons that responded in a manner typical of the recipient species or other neurons that have not previously been found in the type C sensilla of either species. The single-cell data help to explain behavioral results showing that some transplant males do fly upwind to both species' pheromone blends, an outcome not expected based on known antennal sensory phenotypes. Our results suggest that host tissue can influence antennal olfactory receptor neuron development, and further that because of a common phylogenetic ancestry the donor tissue has the genetic capability to produce a variety of sensillar and receptor types.     

Differentiation of insect sensory neurons in the absence of their normal synaptic targets.

Sensory neurons in the antenna of the moth, Manduca sexta, arise and differentiate during the 18 days of metamorphosis from pupa to adult, sending axons to the brain. To assess the trophic dependence of developing antennal neurons on their targets, we studied antennae from surgically debrained animals. If the brain is removed 1 to 45 hr after pupal ecdysis (before and during the period when antennal neurons arise by cell divisions), adult development can be triggered by injection of β-ecdysone; if the brain is removed 50 to 60 hr after pupal ecdysis (before antennal axons contact the brain), metamorphosis proceeds spontaneously. Neurons proliferate normally and differentiate extensively in the antennae of debrained animals. They acquire a characteristic size and shape, elaborate axons and dendrites, migrate to appropriate positions in the sensilla, accumulate components of a neurotransmitter system (acetylcholine, choline acetyltransferase, and acetylcholinesterase), and generate electrical responses to olfactory and mechanical stimuli. Antennal sensory neurons thus differ from a variety of vertebrate neurons, which fail to mature when deprived of their synaptic targets.     

The <Emphasis Type="Italic">Tribolium spineless</Emphasis> ortholog specifies both larval and adult antennal identity

The morphology of insect antennae varies widely among species, but our understanding of antennal development comes almost solely from studies of one species—the fruit fly, Drosophila melanogaster. Moreover, this knowledge applies mostly to adult structures, since Drosophila lacks external larval appendages. In contrast to Drosophila, the red flour beetle, Tribolium castaneum, has both larval and adult antennae, which are very different from one another in morphology. Thus, Tribolium provides an ideal system to compare modes of antennal development both within and between species. Here, we report that the Tribolium ortholog of spineless (Tc-ss) is required in both the larval and adult antennae. Knockdown of Tc-ss by RNAi during either larval or imaginal development causes transformation of the distal portion of the antennae to legs. Thus, the function of ss is conserved between Drosophila and Tribolium with respect to adult antennal specification and also between Tribolium larval and adult antennal development. The similarity of the Tc-ss RNAi phenotype to that of a classically described Tribolium mutation, antennapedia (ap) (of no relationship to the Drosophila Hox gene of the same name), led us to characterize the original ap mutation and two newly identified ap alleles. Our mapping and phenotypic data suggest that Tc-ss is the best candidate for the ap locus. These results represent a first step in characterizing larval and adult antennal patterning in Tribolium, which should provide important insights into the evolution of insect antennal development.     

Expression pattern in the antennae of a newly isolated lepidopteran Gq protein alpha subunit cDNA.

From the antennae of the moth Mamestra brassicae, we have identified a lepidopteran G protein alpha subunit belonging to the Gq family, through immunological detection in crude antennal extract and antennal primary cell cultures, followed by molecular cloning. The complete cDNA sequence (1540 bp) contains an open reading frame encoding a protein of 353 amino acids. This deduced sequence possesses all of the characteristics of the Gq family and shares a very high degree of amino-acid sequence identity with vertebrate (80% with mouse or human Gqalpha) and invertebrate subunits (varying between 60 and 87% for Gqalpha from organisms as diverse as sponge and Drosophila). The expression pattern of the Gq subunit in adult antennae was associated with the olfactory sensilla suggesting a specific role in olfaction. These data provide molecular evidence for a component of the phosphoinositide signaling pathway in moth antennae: this G protein alpha subunit may be involved in the olfaction transduction process through interaction with G-protein-coupled receptors, stimulating the phospholipase C mediated second messenger pathway.     

Antennal and locomotor responses to attractive and aversive odors in the searching cockroach

The behavioral responses to attractive and aversive odors were examined in blinded adult male cockroaches under tethered-walking conditions. A sex pheromone-like stimulant derived from adult virgin females and artificially synthesized limonene were used as attractive and aversive odor sources, respectively. When a searching animal was stimulated with the attractive female-derived odor, the horizontal deflections of both the antennae were increased, and in most cases the vertical antennal positions were shifted downward. The stimulation also significantly decreased the walking speed of the animal. These behavioral changes imply a careful search in the immediate surroundings. The aftereffect of the sex pheromone was more pronounced on locomotion than on antennal movement. On the other hand, stimulation with the aversive odor (limonene) tended to suppress active antennal movement, and also increased the walking speed. Immediately after the withdrawal of the aversive odor, the active movement of the antennae was resumed, and the walking speed rapidly decreased to a level approximately the same as that of the control period. These results indicate that the responses to the qualitatively opposite types of odor are reciprocal to each other with regard to both antennal movement and locomotion.     

Directionality of antennal sweeps elicited by water jet stimulation of the tailfan in the crayfish Procambarus clarkii

Summary Directionality and intensity dependence of antennal sweeps elicited by water jet stimulation of the tailfan in tethered, reversibly blinded adult and juvenile crayfish (Procambarus clarkii) were analyzed.Resting crayfish keep their antennae at about 50° symmetrically to the longitudinal body axis (Figs. 2 bottom, and 3).In adults, tailfan stimulation elicits synchronous backward sweeps of both antennae, which increase for more caudal stimulus directions (Figs. 2–4 and 5A). Directions differing by 30°–60° are significantly distinguished (Fig. 4). The mean sweep of the ipsilateral antenna significantly overrides that of the contralateral antenna for rostrolateral stimulation at 40–200 mm/s stimulus velocity and lateral to caudolateral stimulation at 40 mm/s and thus lateralization of the stimulus is revealed (Figs. 2 top, 4 and 5A). Mean antennal sweeps at a given stimulus direction and distance increase with increasing stimulus velocity (40–250 mm/s, Fig. 5A).In juveniles, the directional dependence of antennal sweeps is reduced compared to that of adults, while a similar intensity dependence is found (Fig. 5B).The pronounced directionality of the antennal response in adult crayfish vanishes and response latencies increase after reversibly covering the tailfan with a small bag or the telson with waterproof paste (Figs. 6 and 7). Thus, tailfan and especially telson mechanoreceptors play an important role in the localization of water movements elicited by predators or prey behind the crayfish.     

External morphology of the antennae of Damalinia ovis (Phthiraptera: Trichodectidae)

The antennae of adult Damalinia ovis, the sheep louse, were studied using light and scanning electron microscopy. Sensory structures are located on all three antennal segments with the predominant sensilla type being tactile. Nine different types of sensilla are described on the basis of external appearance. One of the sensilla, designated a "pit organ" because of its unusual shape, has not been described previously. A pair of these sensilla are present on each antenna, and their function is unknown. A group of 11 sensilla on the tip of each antenna contains olfactory and chemosensory pegs, and a possible thermohygroreceptor. The antennae are sexually dimorphic, the male having more tactile sensilla, two well-developed terminal hooks, and a different cuticular architecture on the posterior surface of antennal segment 1.     

Antennal responses of Cydia pomonella (L.) exposed to surfaces treated with methoxyfenozide

Electroantennogram (EAG) measurements were recorded from the antennae of male and female codling moth, Cydia pomonella L., to determine whether adult moths exposed to surfaces treated with the ecdysteroid agonist methoxyfenozide experience a decline in their antennal reception and thus olfactory sensitivity. Such a phenomenon would offer a possible mechanism for the previously reported decreased responsiveness from moths treated with methoxyfenozide to pheromone‐ and plant volatile‐based monitoring lures. Mean EAG data revealed that the antennae from methoxyfenozide‐treated male moths appear to be just as sensitive to various doses of synthetic codlemone as the antennae from the control and surfactant‐treated moths, but they appeared to be less sensitive to the pheromone component 12OH (collected from female effluvia) than the control male antennae. Mean male EAG responses to the pheromone components E8,E10‐12Al and codlemone collected from methoxyfenozide‐treated females were significantly less than the responses towards those two pheromone components collected from the control and surfactant‐treated females. Female moth exposure to methoxyfenozide did not negatively impact the sensitivity of female antennae to the plant volatile pear ester, but it did towards the apple volatile butyl hexanoate. Data from this study show that adult C. pomonella exposure to methoxyfenozide‐treated surfaces does appear to negatively impact, in a minor way, the (i) olfactory sensitivity (or detection) of male antennae towards some components of the female sex pheromone, (ii) the female antennal sensitivity towards a key apple volatile and (iii) the attractiveness of female pheromone effluvia.     

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.     

Tactile localisation: the function of active antennal movements in the crayfishCherax destructor

Video recordings and single frame analysis were used to study the function of the second antennae of crayfish (Cherax destructor) as a sensory system in freely behaving animals. Walking crayfish move their antennae back and forth through horizontal angles of 100 degrees and more, relative to the body long axis. At rest, animals tend to hold their antennae at angular positions between 20 and 50 degrees. Movements of the two antennae are largely independent of each other. Before and during a turn of the body the ipsilateral antenna is moved into the direction of the turn. Solid objects are explored by repeatedly moving the antennae towards and across them. Both seeing and blinded crayfish can locate stationary objects following antennal contact. On antennal contact with a small novel object, a moving animal withdraws its antenna and attacks the object. When the antenna of a blinded crayfish is lightly touched with a brush the animal turns and attacks the point of stimulation. The direction taken and the distance covered during an attack can be correlated with: the angle at which the antenna is held at the moment of contact and the distance along the antennal flagellum at which the stimulus is applied. From behavioural evidence we conclude that crayfish use information about the angular position of their antennae and about the position of stimulated mechanoreceptors along the antennal flagellum to locate objects in their environment. We suggest ways in which an active tactile system-like the crayfish's antennae--could supply animals with information about the three-dimensional layout of their environment.     

First descriptions of early- and middle-stage copepodids of Anthosoma crassum (Dichelesthiidae: Siphonostomatoida) and lesions on shortfin makos (Isurus oxyrinchus) infected with A. crassum

Early- and middle-stage copepodids of Anthosoma crassum (Dichelesthiidae: Siphonostomatoida) and lesions associated with A. crassum infections are described from samples collected from the jaws of shortfin makos captured off southern California. The copepodids did not possess frontal filaments or frontal organs, and they resided in a headstandlike position firmly attached by their embedded antennae. Copepod larvae and small adults were lodged in shallow mucosal ulcers that basally exhibited mild, acute granulocytic stomatitis; large adults were lodged in deep tunnels encompassing the anterior aspects of their bodies. Some lesions contained more than I copepod. Examinations of lesions revealed that A. crassum infection of shortfin makos can result in severe subacute, necrotizing stomatitis with hemorrhage, granulation tissue, and lymphocytic aggregations in the mucosa, and reactive lymphocytic infiltration of the submucosal skeletal muscle. Copepod gut contents consisted of shark erythrocytes, hemosiderin granules, and necrotic host cells. These observations, along with reports of sharks heavily infected with A. crassum, suggest that this copepod may sometimes play a role in the morbidity and mortality of sharks that it infects.     

Structure and development of the insect antennodeutocerebral system

The structure and postembryonic development of antennae and deutocerebrum in various insect orders are reviewed. First, the number and/or size of system components, i.e. antennal sensilla, neuroreceptors, deutocerebral neurons and synaptic complexes (glomeruli), are compared in adult insects. Second, the neuronal organization of the system is examined. Evidence of projection of the neuroreceptors from the 2 basal antennal segments (scape and pedicel) in the antennal mechanosensory and motor center and from the distal segments (flagellum) in the antennal lobe is discussed. Third, the types of cerebral neurons found in the antennal lobe are described. Fourth, all synaptic contacts between neurons in the antennal lobe take place in discrete glomeruli whose ultrastructure and neurotransmitters are examined. Evidence of individual identifiability of glomeruli is given. Fifth, various sexual dimorphisms present in the antenna and antennal lobe, related or not to sex pheromone perception, are described at sensillar, neuronal, and glomerular levels. Sixth, the postembryonic development of neurons and glomeruli is analyzed in holo- and hemimetabolous insects. In conclusion, the columnar organization of the system is emphasized, alternative models of antennal-neuron projection into glomeruli are considered, and the functional significance of identified glomeruli is discussed.     

Regulation of cyclic GMP elevation in the developing antennal lobe of the sphinx moth,Manduca sexta

In the moth, Manduca sexta, 3′,5′‐guanosine monophosphate (cGMP) is transiently elevated during adult development in about 100 neurons of the antennal lobe. We demonstrate that nearly all of these neurons are local interneurons of the lateral cluster I, that their capacity to show a strong cGMP response during development is regulated by the steroid hormone 20‐hydroxyecdysone, and that in a subpopulation of these neurons cGMP elevation seems to be controlled directly by the gaseous messenger molecule nitric oxide (NO). Treatment with the acetylcholine esterase inhibitor eserine, antennal nerve transection, and electrical stimulation of the antennae suggest that NO/cGMP signaling during development is an activity‐dependent process. Besides input from the antennae, input from the central brain and the ventral ganglia is involved in upregulating cGMP in the antennal‐lobe neurons. Possible sources are centrifugal aminergic neurons, since application of serotonin and histamine enhances the GMP signal in local interneurons. Comparing the time course of cGMP elevation with events occurring during development leads us to the hypothesis that the NO/cGMP signaling pathway might be involved in synapse formation of a subset of antennal‐lobe neurons. © 1999 John Wiley & Sons, Inc. J Neurobiol 41: 359–375, 1999     

The antennal motor system of the stick insect Carausius morosus: anatomy and antennal movement pattern during walking

The stick insect Carausius morosus continuously moves its antennae during locomotion. Active antennal movements may reflect employment of antennae as tactile probes. Therefore, this study treats two basic aspects of the antennal motor system: First, the anatomy of antennal joints, muscles, nerves and motoneurons is described and discussed in comparison with other species. Second, the typical movement pattern of the antennae is analysed, and its spatio-temporal coordination with leg movements described. Each antenna is moved by two single-axis hinge joints. The proximal head-scape joint is controlled by two levator muscles and a three-partite depressor muscle. The distal scape-pedicel joint is controlled by an antagonistic abductor/ adductor pair. Three nerves innervate the antennal musculature, containing axons of 14-17 motoneurons, including one common inhibitor. During walking, the pattern of antennal movement is rhythmic and spatiotemporally coupled with leg movements. The antennal abduction/adduction cycle leads the protraction/retraction cycle of the ipsilateral front leg with a stable phase shift. During one abduction/adduction cycle there are typically two levation/depression cycles, however, with less strict temporal coupling than the horizontal component. Predictions of antennal contacts with square obstacles to occur before leg contacts match behavioural performance, indicating a potential role of active antennal movements in obstacle detection.     

Investigation of the sensory organs on antennae of the horseflies Hybomitra bimaculata and Tabanus bovinus (Diptera: Tabanidae) by scanning electron microscope

Sensory organs on the antennae of the horseflies Hybomitra bimaculata Macq. and Tabanus bovinus Loew are represented by the same morphological types of sensilla. Never differences in the topographical distribution of the sensilla on antennae have been also found, which can be explained by the similarity of ecological and behavioural adaptations of these insects. First and second antennal segments are found to be supplied with tactile hairs and proprioceptors. Other antennal segments bear sensory organs of several morphological types. Short thin olfactory hairs are most numerous among them. They are present on all segments of the antennal flagellum and belong to two morphological types different by the hair length. In the upper parts of the antennal segments from third to seventh several sensilla trichoidea are present, which probably serve as tactile and taste receptors.     

Antennal courtship and functional morphology of tyloids in the parasitoid wasp Syrphoctonus tarsatorius (Hymenoptera: Ichneumonidae: Diplazontinae)

Courtship behaviour and associated morphological characters are believed to evolve under diversifying sexual selection. In Hymenoptera, sexually dimorphic antennal structures, the ‘tyloids’, show a large variability. Although crucial for functional interpretation, the link between tyloid morphology and courtship behaviour has gained only limited attention. Here, we investigate antennal morphology and antennal courtship in the parasitoid wasp Syrphoctonus tarsatorius (Hymenoptera: Ichneumonidae: Diplazontinae). We confirm the glandular nature of the tyloids by light and scanning electron microscopy. Moreover, we report a new form of antennation during courtship, antennal double-coiling, which links morphology and behaviour by bringing the tyloids in direct contact with the antennae of the female, thus probably facilitating the transfer of a contact pheromone. We show that a change in haemolymph pressure is the activator of the antennal movement and that it can be reproduced in the laboratory using amputated antennae. Investigations of antennal structure and movement in three additional hymenopteran species suggest that the number and location of tyloids coincide with the modality of antennal coiling. Our method for simulating antennal movement will enable retrieving information about courtship behaviour from museum specimens, thus leading to a better understanding of the evolution of courtship behaviour in Hymenoptera.     

Antennal sensilla and sexual dimorphism of the adult ladybird beetle Semiadalia undecimnotata Schn. (Coleoptera: Coccinellidae)

The antennal receptors of the adult male and female ladybird beetle, Semiadalia undecimnotata (Coleoptera: Coccinellidae), were examined by scanning and transmission electron microscopy. Twelve types of receptors were characterized and grouped into 5 morphological classes: Böhm, trichoid, coeloconic, basiconic, and chetiform sensilla. Sensory function was determined on the basis of sensillar ultrastructure and electrophysiological response. Olfactory sensilla are confined in both sexes to the 2 terminal antennal segments. In contrast, gustatory and mechanosensitive organs are present along the entire length of the antennae. Sexual dimorphism of antennal receptors is limited to the latter 2 functional classes. The principal characteristics of this dimorphism are the following: a) males possess 540 sensilla (all types), while females possess only 500; b) males exhibit 2 types of taste receptors not present in females; c) females exhibit one type of mechanoreceptor absent in males; d) the 3 sex-specific types of sensilla, which occupy the same position in males and females, are confined to the inner side of the antennae. The possible role of male-specific sensilla in intersexual communication is discussed.