Distal Antennal Segments Research Articles

Detailed morphological structure and phylogenetic relationships of Degeeriella punctifer (Phthiraptera: Philopteridae), a parasite of the bearded vulture Gypaetus barbatus (Accipitriformes: Accipitridae)

Habitat loss is one of the main threats to species survival and, in the case of parasites, it is their hosts that provide their habitat. Therefore, extinction even at local scale of host taxa also implies the extinction of their parasites in a process known as co-extinction. This is the case of the bearded vulture (Gypaetus barbatus), which almost became extinct at the beginning of the twentieth century. After several attempts, this species was successfully reintroduced into the Alps at the end of the twentieth century. We collected 25 lice specimens from an electrocuted bearded vulture from Susa (Italian Alps) that were morphologically identified as Degeeriella punctifer. Six individuals were studied by scanning electron microscopy, with particular emphasis on their cephalic sensorial structures, while four further specimens were characterized at molecular level by amplifying partial regions of the 12SrRNA, COX1 and elongation factor 1 alpha (EF-1) genes. From a morphological perspective, the number, type and arrangement of the sensillae on the two distal antennal segments is quite similar to that of other species of the family Philopteridae (Phthiraptera: Ischnocera). The mandibles and tarsal claws allow lice to cling firmly to their host’s feathers. Phylogenetic analyses help unravel the paraphyletic nature of the genus Degeeriella and demonstrate the clear differentiation between lice parasitizing Accipitriformes and Falconiformes, as well as the close relationship between D. punctifer, D. fulva, D. nisus and Capraiella sp. that, along with other genera, parasitize rollers (Aves: Coraciiformes).

Electroantennographic Responses of Aromia bungii (Faldermann, 1835) (Coleoptera, Cerambycidae) to a Range of Volatile Compounds.

Background: The red-necked longhorn beetle, Aromia bungii, is one of the most damaging pests of stone fruit trees. Native to the south-eastern Palearctic and Oriental regions, it invaded and is established to some extent in the Campania Region (Southern Italy). In several cerambycid species, volatile organic compounds (VOCs) have been shown to play a role in mate and host plant location. Methods: The electroantennographic (EAG) technique was employed to explore the antennal chemoreceptivity of male and female A. bungii antennae to 90 VOCs. Results: Increasing EAG amplitudes from the basal to the distal antennal segments were recorded in response to six selected plant volatiles. From the distal flagellomeres, the largest EAG responses (>0.8 mV) were elicited by 2-hexanol, octanal, sulcatone, guaiacol, sulcatol, 2,4-dimethyl-3-hexanol, 2,4-dimethyl-2-hexanone, heptanal, nonanal, (Z)-3-hexenol, and 1-heptanol in both sexes, and by linalool, (E)-2-heptenal, 1-octen-3-ol, (E)-2-octenal, 3-octanol, (E)-2-octen-1-ol, α-phellandrene, and α-terpinene in males. The olfactory system of both sexes proved to be sensitive to changes in stimulus concentration and compound structure. Conclusions: this study demonstrates the capability of A. bungii males and females to detect and discriminate among a wide range of VOCs and provides a basis for further olfactometer and field trapping experiments aimed at identifying behaviorally-active compounds useful for the implementation of semiochemical-based control strategies for this pest.

Transcription factor Dlx3 induces aryl hydrocarbon receptor promoter activity

The Distal-less (Dlx) homeobox transcription factors (TFs) play a prominent role in regulating multiple facets of vertebrate biology. Though widely studied as mediators of tissue development, recent work has uncovered a role for this TF family in modulating the vertebrate hematopoietic compartment. Pertinent to our study, murine Dlx1-3 are expressed in an innate lymphocyte population known as natural killer (NK) cells, and they are implicated to assume a functional role in the NK cell maturation pathway. However, Dlx target genes are poorly understood. In Drosophila, the invertebrate Dlx ortholog Distal-less (Dll) regulates another transcription factor called Spineless (ss), which is critical for specifying distal antennal segments. Importantly, the vertebrate ortholog of ss is the aryl hydrocarbon receptor (AhR), a transcription factor recently shown to be important in the regulation of a number of immune cell subsets, including NK cells. Given these findings, we investigated whether Dlx TF family members might analogously regulate AhR in an NK cell context. Our results demonstrate that Dlx3 is constitutively co-expressed with AhR in murine and human CD127+ NK cells. Critically, we show that Dlx3 induces AhR promoter activity by binding to a regulatory region that resides ~5.5kb upstream of the transcriptional start site. This mechanism is functionally relevant, as Dlx3 expression in human NK cells significantly enhances TF activity at AhR DNA-binding elements (Xenobiotic Responsive Elements, XREs). Thus, our study defines Dlx3 as a positive regulator of the aryl hydrocarbon receptor.

Antennal cropping during colony foundation in termites

The literature on pairing and mating behavior in termites indicates that a number of distal antennal segments in dealates of both sexes are often removed during colony foundation, with terms such as amputation, mutilation and cannibalism typically employed to report the phenomenon. Here we propose the use of the phrase ‘antennal cropping’ to describe the behavior, and assess naturally occurring levels of its occurrence by comparing the number of antennal segments in museum specimens of alates and dealates in 16 species of Australian termites (four families), supplemented by analyzing published data on Coptotermes gestroi. Dealates had significantly fewer antennal segments than alates in 14 of the 16 termite species, with both exceptions belonging to the family Termitidae. Levels of antennal cropping were not significantly different between the sexes but did vary by family. Dealates in the Kalotermitidae removed the most segments (41.3%) and those in the Termitidae removed the fewest (8.9%). We discuss the biological significance of this phylogenetically widespread termite behavior, and suggest that controlled antennal cropping is not only a normal part of their behavioral repertoire but also a key influence that changes the conduct and physiology of the royal pair during the initial stages of colony foundation.

Mixture Interactions in Moth Olfactory Physiology: Examining the Effects of Odorant Mixture, Concentration, Distal Stimulation, and Antennal Nerve Transection on Sensillar Responses

The insect olfactory system is challenged to decipher valid signals from among an assortment of chemical cues present in the airborne environment. In the moth, Heliothis virescens, males rely upon detection and discrimination of a unique blend of components in the female sex pheromone to locate mates. The effect of variable odor mixtures was used to examine physiological responses from neurons within sensilla on the moth antenna sensitive to female sex pheromone components. Increasing concentrations of heliothine sex pheromone components applied in concert with the cognate stimulus for each neuronal type resulted in mixture suppression of activity, except for one odorant combination where mixture enhancement was apparent. Olfactory receptor neuron (ORN) responses were compared between moths with intact and transected antennal nerves to determine whether specific instances of suppression might be influenced by central mechanisms. Type A sensilla showed little variation in response between transected and intact preparations; however, recordings from type B sensilla with transected antennal nerves exhibited reduced mixture suppression. Testing by parallel stimulation of distal antennal segments while recording and stimulating proximal segments dismissed the possibility of interneuronal or ephaptic effects upon sensillar responses. The results indicate that increasing concentrations of "noncognate" odorants in an odor mixture or antennal nerve transection can produce variation in the intensity and temporal dynamics of physiological recordings from H. virescens ORNs.

Off-host aggregation behavior and sensory basis of arrestment by Cimex lectularius (Heteroptera: Cimicidae)

The bed bug, Cimex lectularius, aggregates under filter paper disks previously stained by adults. A multiple-choice assay was used to determine differences in aggregation behavior among two strains, multiple lifestages, and levels of starvation. There were no differences in level of aggregation between established and recently derived strains, or among adults and nymphs of different instars. Propensity to aggregate decreased with time since feeding, but preference for stained disks remained high. We also examined which sensory structures mediate aggregation, and whether antennectomy affected movement, orientation, and arrestment under stained disks. Bed bugs that were left intact, blinded, or surgically altered by the removal of probosci or the distal antennal segments exhibited high levels of aggregation under stained disks. However, the removal of the pedicel significantly reduced aggregation compared to intact bugs. Video recordings of movement and orientation by bugs with intact, partial and complete antennectomies demonstrated that neither partial nor complete antennectomies affected walking speed, path straightness, direction of movement or frequency of encounters with either stained or clean disks. However, complete removal of both antennae significantly reduced the percentage of encounters with stained disks that resulted in arrestment. These findings suggest aggregation by bed bugs is a result of arrestment mediated by direct, close-range contact between sensilla on the pedicel and stained disks.

Regulation of the Drosophila distal antennal determinant spineless

The transformation of antenna to leg is a classical model for understanding segmental fate decisions in Drosophila. The spineless ( ss) gene encodes a bHLH-PAS transcription factor that plays a key role in specifying the identity of distal antennal segments. In this report, we identify the antennal disc enhancer of ss and then use enhancer- lacZ reporters to work out how ss antennal expression is regulated. The antennal determinants Distal-less ( Dll) and homothorax ( hth) are key activators of the antennal enhancer. Dll is required continuously and, when present at elevated levels, can activate the enhancer in regions devoid of hth expression. In contrast, homothorax ( hth) is required only transiently both for activation of the enhancer and for specification of the aristal portion of the antenna. The antennal enhancer is repressed by cut, which determines its proximal limit of expression, and by ectopic Antennapedia (Antp). Repression by Antp is not mediated by hth, suggesting that ss may be a direct target of Antp. Finally, we show that ss + is not a purely passive target of its regulators: ss + partially represses hth in the third antennal segment and lies upstream of Dll in the development of the maxillary palp primordia.

Temporally Staggered Glomerulus Development in the Moth Manduca sexta

Glomeruli, neuropilar structures composed of olfactory receptor neuron (ORN) axon terminals and central neuron dendrites, are a common feature of olfactory systems. Typically, ORN axons segregate into glomeruli based on odor specificity, making glomeruli the basic unit for initial processing of odorant information. Developmentally, glomeruli arise from protoglomeruli, loose clusters of ORN axons that gradually synapse onto dendrites. Previous work in the moth Manduca sexta demonstrated that protoglomeruli develop in a wave across the antennal lobe (AL) during stage 5 of the 18 stages of metamorphic adult development. However, ORN axons from the distal segments of the antenna arrive at the AL for several more days. We report that protoglomeruli present at stage 5 account for only approximately two or three of adult glomeruli with the number of structures increasing over subsequent stages. How do these later arriving axons incorporate into glomeruli? Examining the dendritic projections of a unique serotonin-containing neuron into glomeruli at later stages revealed glomeruli with immature dendritic arbors intermingled among more mature glomeruli. Labeling ORN axons that originate in proximal segments of the antenna suggested that early-arriving axons target a limited number of glomeruli. We conclude that AL glomeruli form over an extended time period, possibly as a result of ORNs expressing new odorant receptors arriving from distal antennal segments.

External morphology of antennal and rostral sensillae in four hemipteran insects and their possible role in host plant selection

The sensory receptors on the antennal flagellar segments and on the rostral tip of four adult hemipteran insects were identified and compared using scanning electron microscopy in order to better understand the role of sensillae in the process of host plant selection and the structural adaptations of feeding organs for phytophagy. The species studied are Riptortus pedestris Fabricius (Alydidae), Elasmolomus sordidus (Fabricius) (Lygaeidae), Cyclopelta siccifolia Westwood (Pentatomidae) and Chrysocoris purpurea (Westwood) (Pentatomidae). Differences in the distribution and arrangement of sensory receptors in the hemipteran families were noted, especially in the shape of the antennal apices, which are of taxonomical significance. A large array of trichoid basiconic and coeloconic sensillae is distributed on the antennal surface. Distal antennal segments have more sensillae and more types of sensillae than the proximal segments, indicating their prominent role in host plant surface exploration. There is only a slight variation in the shape of the stylets owing to the similarity in feeding habits. The mandibular tips are straight and sharp and do not contain any hooks or barbs as in carnivorous species.

Muscular anatomy and aspects of the structure of the antennae of Anthosoma crassum (Dichelesthiidae: Siphonostomatoidae: Copepoda).

The muscular anatomy and fine structure of the antennae of adult Anthosoma crassum are described from specimens collected from jaws of shortfin makos. Casual observations of live parasites showed that embedded adult A. crassum can move toward or away from the host by shortening or lengthening the antennae, respectively. These movements were controlled by antennal retractor muscles. Microprotuberances located on the distal antennal segments may increase friction between these copepods and their hosts, thus strengthening parasite attachment.

Limb type-specific regulation of bric a brac contributes to morphological diversity.

The insect antenna and leg are considered homologous structures, likely to have arisen via duplication and divergence from an ancestral limb. Consistent with this, the antenna and leg are derived from primordia with similar developmental potentials. Nonetheless, the adult structures differ in both form and function. In Drosophila, one conspicuous morphological difference is that the antenna has fewer distal segments than the leg. We propose that this is due in part to the variations in the regulation of bric a brac. bric a brac is required for joint formation, and loss of bric a brac function leads to fusion of distal antennal and leg segments, resulting in fewer total segments. Here, we address how bric a brac is regulated to generate the mature expression patterns of two concentric rings in the antenna versus four concentric rings in the leg. We find that bric a brac expression is activated early throughout most of the Distal-less domain in both antenna and leg and subsequently is restricted to the distal portion and into rings. Although bric a brac expression in the antenna and in all four tarsal rings of the leg requires Distal-less, only the proximal three tarsal rings are Spineless-dependent. Thus bric a brac is regulated differentially even within a single appendage type. The restriction of bric a brac expression to the distal portion of the Distal-less domain is a consequence of negative regulation by distinct sets of genes in different limb types. In the leg, the proximal boundary of bric a brac is established by the medial-patterning gene dachshund, but dachshund alone is insufficient to repress bric a brac, and the expression of the two genes overlaps. In the antenna, the proximal boundary of bric a brac is established by an antenna-specifying gene, homothorax, in conjunction with dachshund and spalt, and there is much less overlap between the bric a brac and the dachshund domains. Thus tissue-specific expression of other patterning genes that differentially repress bric a brac accounts for antenna-leg differences in bric a brac pattern. We propose that the limb type-specific variations in expression of bric a brac repressors contribute to morphological variations by controlling distal limb segment number.

Morphology of Sensory Structures on the Antennae, Maxillary and Labial Palpi, Tarsi, and Ovipositor of Adult Callosobruchus maculatus and C. subinnotatus (Coleoptera: Bruchidae)

The sensilla on the antennae, maxillary and labial palpi, tarsi and ovipositor of adult Callosobruchus maculatus (F.) and C. subinnotatus (Pic) were examined using light and scanning electron microscopy. Six types of antennal sensilla occur in both species. Type 1 are long and have longitudinal ridges and a conspicuous basal socket. These sensilla occur in rings at the distal margin of each antennal segment; the terminal segment bears 2 such rings. Type 2 sensilla are found on all segments and have a well developed basal socket and less conspicuous cuticular ridges. Type 3 sensilla are numerous and found only on flagellar segments. Type 4 sensilla are medium in length and stout. Types 5 and 6 sensilla are short, few, and are found along with type 4 sensilla in groups distally on the distal antennal segments. Types 1 and 2 are sensilla chaetica, type 3 are trichodea, and types 4, 5, and 6 are sensilla basiconica. Sexual dimorphism in numbers of antennal sensilla is observed. Maxillary and labial palps in both species have 3 types of sensilla. Two subtypes in type 3, fluted (3a) and blunt (3b), sensilla are observed in females of both species. Tarsi and ovipositor of both species have 3 types of sensilla each.

Expanded functions in the apical cell domain to regulate the growth rate of imaginal discs

The Drosophila expanded (ex) gene encodes a product (Ex) that shares homology with the Protein 4.1 family of proteins, many of which are enriched at specific lateral cell junctions and the apical cellular domain. Ex colocalizes with actin in the apical domain of imaginal disc epithelial cells, where it partially overlaps the distribution of phosphotyrosine (PY)-containing proteins. This suggests that Ex is present in or associated with adherens junctions. Genetic studies show that Ex is necessary for proper regulation of final cell number in adult wings and for the formation of eyes, distal leg, and distal antennal segments. We have generated mitotic clones that lack Ex using the twin spot technique, and demonstrated that the primary function of Ex is to regulate cell proliferation. Overexpressing Ex protein results in a decrease in final cell number in wings, suggesting a direct relationship between Ex function and proliferation rate. Dev. Genet. 20:103–110, 1997. © 1997 Wiley-Liss, Inc.

Expanded functions in the apical cell domain to regulate the growth rate of imaginal discs.

The Drosophila expanded (ex) gene encodes a product (Ex) that shares homology with the Protein 4.1 family of proteins, many of which are enriched at specific lateral cell junctions and the apical cellular domain. Ex colocalizes with actin in the apical domain of imaginal disc epithelial cells, where it partially overlaps the distribution of phosphotyrosine (PY)-containing proteins. This suggests that Ex is present in or associated with adherens junctions. Genetic studies show that Ex is necessary for proper regulation of final cell number in adult wings and for the formation of eyes, distal leg, and distal antennal segments. We have generated mitotic clones that lack Ex using the twin spot technique, and demonstrated that the primary function of Ex is to regulate cell proliferation. Overexpressing Ex protein results in a decrease in final cell number in wings, suggesting a direct relationship between Ex function and proliferation rate.

Responses and spike generation in the largest antennal taste hairs of Periplaneta brunnea Burm.

The largest taste hair type of the 20 distal antennal segments is investigated electrophysiologically. Two chemoreceptors respond to potassium chloride, tripotassium citrate but not to choline salts, another one to sucrose, but not to D-glucose and only slightly to D-fructose. No response was obtained with pure water. Spikes with a leading negative phase indicate that the primary process controls an apical spike generator localized in the membranes of the outer dendritic segment. The succeeding basal spike consists of two positive phases, that are interpreted as the result of two further spike generation zones, one at the dendritic inner segments and the other at the origin of the axon. Spike activity in the mechanoreceptor also starts with negative going spikes, that are followed by a series of solely positively going ones. TTX applied to the hair tip inhibits the apical spike generator of the chemo- and mechanoreceptor cells.

Fine structure of the coniform sensillar complex on the antennal flagellum of the stonefly nymph Paragnetina media (Plecoptera: Perlidae)

Four to seven coniform sensillar complexes, each formed by 16–21 narrow spines, project perpendicularly from the distal end of middle and distal antennal flagellar segments of the stonefly nymph Paragnetina media. Each spine in the complex is innervated by two neurons with unbranched dendrites. The dendrites extend to a pear-shaped cavity in the tip of the spine. The tip of the spine is porous and contains channels filled with an amorphous material. Each pair of neurons is surrounded by three accessory cells. Based on structural similarities with the terminal multiporous sensilla of other insects, each spine in a sensillar complex represents a fundamental type of chemosensillum.

Internal proprioceptive organs of the distal antennal segments in Allacma fusca (L.) (Collembola : Sminthuridae): Proprioceptors phylogenetically derived from sensillum-bound exteroceptors

The internal proprioceptive organs of the distal antennal segments of Allacma fusca (Collembola : Sminthuridae) were examined with light, scanning, and transmission electron microscopy. There are 2 such organs, each belonging to the sensory equipment of a sensory peg. These pegs are part of the sensory organs of the 3rd antennal segment (SO - AIII), and show characteristics of both hair sensilla and chordotonal organs. They have 5 sensory cells of which 4 can be regarded as chemoreceptors, because the walls of the pegs are pierced by pores. The 5th cells are the proprioceptors. Their dendritic outer segments are connected to the base of the SO - AIII peg but show no specializations. Their dendritic inner segments are distally joined to a thecogen cell containing a scolopale. Proximally, they span the hemolymph space within the antenna. The perikaryon of one of the organs is located on a muscle that deflects the 3rd antennal segment (IP - AIII). The perikaryon of the other is attached to the antennal nerve near a point where it is connected to a muscle that deflects the 4th antennal segment (IP - AIV). There is a conspicuous membrane system, associated with microtubules and a prominent ciliary rootlet within the dendritic inner segment of the IP - AIII. The cytological features of the organs are discussed with reference to 2 problems, the phylogeny of chordotonal organs and the process of mechanosensory transduction.

Antennal Sensilla Coeloconica of Pedicinus obtusus obtusus (Anoplura: Hoplopleuridae)

Recently (Miller 1969; 1970a,b; 1971a) I have shown the sensilla on the distal antennal segments of the Anoplura, previously described as ring sensoria by Ferris (1951), to be either sensilla basiconica or sensilla coeloconica. It was demonstrated also (Miller 1971a) that the size of the openings of the sensilla coeloconica and the distinctness and location of the 2 pore organs found near the distal sensilla vary sufficiently within a genus to be of possible taxonomic value. I (Miller 1969, 1971b) also demonstrated that the number and size of hairs projecting from the apex of the peg of the sensilla basiconica vary from one group to another even within the same families. Similar variations are found among the pegs of the sensilla coeloconica. These findings are not unique to the Anoplura; Clay (1970) demonstrated similar variations in the sensilla coeloconica of the Amblycera.