Ultrastructure of antennal and posterior abdominal sensilla in Chlorophorus caragana females

Chlorophorus caragana Xie & Wang is a destructive wood-boring beetle that damages Caragana spp. bushes and is distributed in desert areas in north-west China. Using scanning electron microscopy and transmission electron microscopy, we observed the morphology and ultrastructure of antennal and posterior abdominal sensilla in C. caragana females, to discuss the putative functions of these sensilla in host location and oviposition behaviors.In total, seven types (24 subtypes) of sensilla were located on the antenna and posterior abdomen. On the antenna, there were Böhm's bristles (BB.); four subtypes of sensilla chaetica (Ch.1–Ch.4) characterized by non-porous surfaces and sensillum-lymph cavities without dendrites; two subtypes of sensilla trichodea (Tr.1 and Tr.2) with a tip pore and dendrites surrounded by dendritic sheaths; dome-shaped sensilla (Dom.) emerging from a deep cavity with one tip pore; four subtypes of sensilla basiconica (Ba.1–Ba.4) and one type of sensilla auricillica (Au.) with a porous cuticular surface and dendrites in the sensillum-lymph cavity; and one type of sensilla styloconica (Sty.) with grooves on the cuticular wall. On the posterior abdomen, there were four subtypes of sensilla chaetica (Ch.5–Ch.8); three subtypes of sensilla trichodea (Tr.3–Tr.5); and three subtypes of sensilla basiconica (Ba.5–Ba.7; Ba.5 had no groove in the cuticular wall, Ba.6 had one tip pore, and Ba.7 was located in a cuticular cavity). The antennal sensilla were believed to be mechanosensitive, chemosensitive, and sensitive to humidity and temperature, and to play roles in mating, host location and oviposition. The abdominal sensilla are believed to be related to oviposition behaviors.     

Antennal morphology and sensillar ultrastructure of Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae)

The parasitoid beetle Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae) is an important parasite of longicorn beetles (Cerambycidae), and has been used in China for the biological control of the Asian longicorn beetle and the Japanese pine sawyer. In this study the antennal morphology and sensillar ultrastructure of D. helophoroides were observed using scanning electron microscopy and transmission electron microscopy. Two types of sensilla trichodea (Tr. 1 and Tr. 2), two types of sensilla basiconica (Ba. 1 and Ba. 2), three types of sensilla chaetica (Ch. 1, Ch. 2 and Ch. 3), and Bohm's bristles were identified according to the morphology and fine structure of each type of sensilla in both sexes. Ultrastructural studies revealed porous structures on the cuticle wall and dendritic branches in the inner lumen of Tr. 1, Tr. 2, Ba. 1, and Ba. 2, thereby suggesting chemoreceptor functions. No difference in shape, structure, sensilla distribution and typology was observed between the sexes. These structures likely have roles in the host locating and habitat searching behavior of adult D. helophoroides, and suggest future studies on the olfaction and host location behavior of D. helophoroides and other coleopteran parasitoids.     

Antenna morphology and sensilla ultrastructure of Tetrigus lewisi Candèze (Coleoptera: Elateridae)

We used scanning and transmission electron microscopy to study the typology, morphology, distributions, and ultrastructures of the antennal sensilla of Tetrigus lewisi Candèze, a predatory click beetle that feeds on longhorned beetles, such as, Monochamus alternatus (Coleoptera: Cerambycidae). We observed eight types of sensilla on the antennae, including sensilla chaetica (with three subtypes: ch.1, ch.2, ch.3), sensilla basiconica (subtypes: ba.1, ba.2, ba.3), sensilla trichodea (subtypes: tr.1, tr.2), as well as sensilla auricillica, sensilla coeloconica, sensilla campaniformia, sensilla styloconica and Böhm's bristles. Significant sexual dimorphism was found in the antenna morphology, as well as in the density of type 2 sensilla trichodea and type 1 sensilla basiconica. We observed thick cuticular walls on sensilla chaetica, sensilla trichodea and sensilla campaniformia; clear pore structures on sensilla trichodea, sensilla basiconica and sensilla auricillica; and double walls with spoke-channels on sensilla coeloconica. The chemoreception, mechanoreception and thermo-/hygro-reception functions were deduced from fine structures on the cuticular walls and the dendrites of the different sensilla types. We suggest that all these sensilla have important roles in the host location, mating and predatory behavior of T. lewisi.     

Antennal morphology and sensilla ultrastructure of Dendroctonus valens LeConte (Coleoptera: Curculionidae,Seolytinae), an invasive forest pest in China

Dendroctonus valens LeConte, an invasive forest pest, is highly dependent on its olfactory system for activities such as host seeking, mating, oviposition and population aggregation. The antennae are the primary olfactory organs in bark beetles. We describe four morphological types of sensilla on the antenna of D. valens: (1) 3 subtypes of sensilla trichodea located at the apex and lateral surface of the club. Sensilla trichodea types 1 and 3 were innervated by 5 and 13 dendrites, respectively. Sensilla trichode type 2 was not found dendrites in a lumen, (2) sensilla chaetica found on the scape and funicle and innervated by 7 dendrites, (3) a pair of sensilla basiconica distributed primarily on the three sensory bands, each innervated by 2 dendrites at the base, and (4) one fluted cone type of grooved peg dispersed throughout the sensory bands. The possible functions of these sensilla are discussed in relation to their morphology and ultrastructure.     

Antennal sensilla of the ground beetle Bembidion properans Steph. (Coleoptera, Carabidae)

The arrangement of antennal sensilla was studied in female and male ground beetles Bembidion properans Steph. (Coleoptera, Carabidae) using scanning electron microscopy. The filiform antennae, 1.8-1.9 mm in length, consist of the scape, pedicel and nine flagellomeres. In both sexes, three types of sensilla chaetica, two types of sensilla trichodea, six types of sensilla basiconica, one type of sensilla coeloconica and one type of sensilla campaniformia were distinguished. The possible function of the sensilla is discussed and three types of sensilla are considered olfactory, sensilla trichodea type 2 and sensilla basiconica types 1 and 2. Olfactory sensilla form dorsal and/or ventral sensillar fields on the flagellomeres and occur sparsely or not at all outside these areas. No sexual differences in the types, number and distribution of antennal sensilla were found.     

Ultrastructure of antennal sensilla of an autoparasitoid Encarsia sophia (Hymenoptera: Aphelinidae)

Encarsia sophia (Hymenoptera: Aphelinidae) is a parasitoid utilized for biological control of Bemisia tabaci, with selection of prey aided by chemoreceptor organs. The morphology and distribution of the antennal sensilla (chemoreceptors) of E. sophia were examined using Transmission electron micrographs. The total antennal length for E. sophia was 429.28 ± 0.95 μm for females and 437.19 ± 8.21 for males, and each antennae was found to consist of seven sensilla of different types. Both sexes possessed sensilla chaetica, sensilla trichodea, basiconic capitate peg sensilla, multiporous grooved-surface placoid sensilla (MG-PS), uniporous rod-like sensilla, nonporous finger-like sensilla, and sensilla coeloconica. Transmission electron micrographs of longitudinal sections of female antennae showed that they were composed of fat body, cuticle, mesoscutello-metanotal muscles, neurons, and glandular tissue, and cross-sections of the basal MG-PS showed sensillar lymph cavities and dendrites. The MG-PSs were imbedded in an electron-dense mass with cuticular invaginations which acted as pores that connected to a central lumen. The possible function of each type of sensilla is discussed.     

Scanning electron microscopy studies of antennal sensilla of bruchid beetles,Callosobruchus chinensis (L.) and Callosobruchus maculatus (F.) (Coleoptera: Bruchidae)

The bruchid beetle, Callosobruchus chinensis (L.) and C. maculatus (F.) (Coleoptera: Bruchidae), are important stored-product insects of stored legume seeds. In this study, the external morphologies of the antennal sensilla on the antennae of both female and male adults of these two species were described using scanning electron microscopy. Antennae of both species are made up of the scape, pedicel, and nine segments of flagellomeres. Antennae of female and male C. maculatus and female of C. chinensis are serrate in shape, while those of male C. chinensis are pectinate. Eight morphological sensilla types were recorded in both sexes, including Böhm bristles (BB), two types of sensilla trichoid (ST1, ST2), sensilla chaetica (SC), two types of sensilla basiconic (SB1, SB2), grooved pegs (GP), and sensilla cavity (SCa). The number of ST1 and SB1 of the male were significantly greater than those of the female of C. chinensis, and the number of ST2 and SB1 of the male were significantly more abundant than those of the female of C. maculatus. The possible functions of the above sensilla types are discussed in light of previously published literature.     

Morphology,distribution and abundance of antennal sensilla in three stink bug species (Hemiptera: Pentatomidae)

The neotropical stink bugs, Euschistus heros, Piezodorus guildinii and Edessa meditabunda, are important pests of soybean and other crops throughout Central America and in South America from Northern Argentina to Brazil. Mate finding and host plant location in these species depend largely on their chemical communication, and semiochemicals are important mediators of these behaviors. In this study scanning electron microscopy was used to examine the external morphology, distribution and abundance of antennal sensilla on males, females and 5th instar nymphs of these species. Nine morphologically different sensilla types were found: trichod sensilla, type 1 and 2 (ST1 and ST2), long and short basiconic sensilla (SB1, SB2, and SB3), slit-tipped and knob-shaped basiconic sensilla, long chaetic sensilla (Sch) and coeloconic sensilla (Sco). Differences were detected in the abundance and arrangement of the sensilla over the antennal segments in individuals of the same species and among the species studied. The Sch, Sco and the slit-tipped and knob-shaped basiconic sensilla accounted for the major difference in sensilla types among the species. The ST1 was the most abundant type and was restricted to the flagellum. The pedicel of E. heros differs from the pedicels of P. guildinii and E. meditabunda mainly by the absence of Sch. There was a sexual dimorphism in ST1, SB1 and SB2, and this may be an indicative of their roles in detection of male-produced sex pheromone and odors derived from the host plants. The SB2 was lacking in the antennal tip of both sexes and 5th instar nymphs, but was abundant on the second flagellar segment of females of the three species. The same types of sensilla were found on 5th instar nymphs, but always in significantly lower numbers. The morphology and putative functions of each sensilla were compared and discussed.     

Antennal sensory receptors of Pteromalus puparum female (Hymenoptera: Pteromalidae), a gregarious pupal endoparasitoid of Pieris rapae

The external morphology of the antennal sensilla of Pteromalus puparum females (Hymenoptera: Pteromalidae) is described using scanning electron microscopy. The antennae of P. puparum females are geniculate in shape, formed from a large, cylindrical scape with a basal radicel fitting into the antennal socket, a shorter, barrel-shaped pedicel and a flagellum composed of 12 subsegments. Eight morphologically distinct types of sensilla were found on the female antennae. These are: nonporous sensilla trichodea types 1 and 2, which are putative mechanosensilla, nonporous sensilla chaetica, which may function as proprioceptors, uniporous sensilla trichodea types 3 and 4, which are presumably contact chemosensilla, basiconic capitate peg sensilla, which probably function in thermo-hygro reception, multiporous sensilla trichodea, and multiporous sensilla placodea which are all presumed to be olfactory sensilla.     

Morphological characterization of the antennal sensilla of the Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae), with reference to their probable functions

The Asian citrus psyllid, Diaphorina citri Kuwayama (Hemiptera: Psyllidae), is one of the most significant economic pests of citrus worldwide. This insect vectors three phloem-restricted bacteria in the genus Candidatus Liberibacter that cause huanglongbing (citrus greening), the most severe disease limiting citrus production worldwide. We examined the external morphology of the antennal sensilla of male and female D. citri using scanning electron microscopy (SEM) and determined the putative functions of the identified sensilla using transmission electron microscopic (TEM) techniques. The filiform antennae of D. citri were of the conventional type comprised of a basal scape, pedicel and a long, thread-like flagellum, which is composed of eight flagellomeres. Eleven morphologically unique sensillar types were found and described on the antennae of male and female D. citri. Of those identified, the two apical setae, multiporous types I and II sensilla trichoidea, and the antennal rhinaria were porous and may be involved in perception of host- and mate-related volatile chemicals. However, the aporous types I, II and III sensilla trichoidea may have mechanosensory functions and the chaetica sensilla, cavity sensilla and unidentified uniporous sensilla may be involved in proprioception, thermo-hygroreception and cold detection, respectively. The shape, external morphology and array of sensilla on the antennae of male and female D. citri were similar. The only major difference observed was in the morphology of the short apical setae, whose tips were recessed inward in females but not so in males. The results are discussed in relation to plausible roles of the identified sensilla in mate and host location by this species.     

Sensilla on the antennal funiculus of the blow fly, Protophormia terraenovae (Diptera: Calliphoridae)

The morphology of the antennal funiculus and the external morphological characteristics and distribution of sensilla of blow fly, Protophormia terraenovae, have been studied using light and scanning electron microscopy. Cross section of the funiculus is roughly triangular in shape, with an anterior-medial, anterior-lateral, and posterior surface. The latter presents some large-size pits on restricted lateral and median areas of the proximal funiculus, and several smaller-size ones close to the pedicel-funiculus joint. The entire surface of the antennal sub-segment appears densely populated by microtrichia and is inhabited by seven types of sensilla: one trichoid, two basiconic, one auriculate, one coeloconic, and two basiconic-like pit sensilla. Trichoid, basiconic, auriculate and basiconic-like types display a multiporous wall, a feature characteristic of insect olfactory sensilla. It remains to be verified whether or not the coeloconic structure type has wall pores. The most abundant sensilla are the trichoid ones, which are followed by the basiconic, coeloconic and auriculate types in a decreasing density order. The basiconic-like pit sensilla are present only on the posterior funicular surface, unlike the remaining ones which populate the entire sub-segment. The blow fly' funiculus displays a significant, even though moderate sexual dimorphism, the female sub-segment being bigger and presenting a higher number of trichoid and auriculate sensilla. The presence of multiple wall pores in most of sensilla types implies an olfactory modality for sensory neurons they accomodate, thus indicating that the blow fly' funiculus is a plain olfactory organ.     

Structure and distribution of antennal sensilla of the red imported fire ant

The morphology of the antenna of the red imported fire ant, Solenopsis invicta, was examined by light microscopy, scanning electron microscopy, and transmission electron microscopy. The antennae are sexually dimorphic: the worker antenna has porous sensilla on the two distal segments (the antennal club), whereas the clubless male antenna has porous sensilla on all segments past the pedicel. The major type of porous sensilla on both male and female is sensilla tricodea curvata. However, the male s. tricodea curvata are rather uniform in size, whereas the female s. tricodea curvata vary considerably in thickness. The number of sensilla on the distal segment of the worker antenna increases with segment length. This suggests a possible mechanism by which task assignments in S. invicta could be determined by the presence or absence of sensilla sensitive to specific task-related odor or pheromone cues. The sensilla basiconica have an invariant spatial pattern on worker and queen antennae.     

Distribution and morphometric studies of flagellar sensilla in Emphorini bees (Hymenoptera, Apoidea)

The tribe Emphorini is a group of pollen-collecting solitary bees with a geographical distribution restricted to the western hemisphere. Most of the Emphorini bees collect Page 10 linepollen from a few specific plant families and display specialized behaviors for constructing their nests. Insect sensilla are the basic structural and functional units of cuticle receptors, serving mainly mechano- and chemo-receptor functions. The external morphology of the antennal sensilla has been well characterized in species of different families of Apoidea, however there is scarce information about this issue in solitary bees of the family Apidae. For a better understanding of the association between the external sensory system and several types of behaviors which emerged along the evolutionary history of bees, it is important to characterize the antennal receptors in several representative species of this tribe. The distribution of the antennal sensilla on the dorsal flagella of 18 taxa was studied in insects of both sexes, using light and scanning electron microscopy. There were six types of sensilla and setae on the antennae, which were identified as sensilla placodea, trichodea, basiconica, coeloconica, coelocapitular and ampullacea. The sensilla trichodea were classified into subtypes, A, B, C-D. Sensilla subtype A were the most abundant sensilla and were distributed over the entire antennae, while sensilla placodea and sensilla trichodea type B, showed a restricted distribution on specific areas of the flagella. We have recognized four patterns of spatial distribution of setae on dorsal flagella. Species having setae on the distal part of the flagellomeres tended to contain a low density of sensilla trichodea type A. Females showed a higher number of sensilla subtypes B and C-D than males; instead sensilla trichodea A were more abundant in males. No significant difference was found in the number of sensilla placodea, ampullacea, coeloconica and coelocapitular. Sensilla basiconica were found only in females. Our results showed that gustative and tactile sensilla were more abundant in female bees, as well as, olfactory receptors predominate in the antennal system of males. The possible coevolution of flagellar sensilla in males and females of solitary bees is discussed in light of previous reports. Patterns of distribution of setae determine the relative abundance of the types of sensilla in the flagellum.     

Antennal sensilla of some forensically important flies in families Calliphoridae, Sarcophagidae and Muscidae

Antennal sensilla of some forensically important fly species in the families Calliphoridae (Chrysomya megacephala, Chrysomya rufifacies, Chrysomya nigripes and Lucilia cuprina), Sarcophagidae (Parasarcophaga dux) and Muscidae (Musca domestica) were studied using scanning electron microscopy. Five types of sensilla were observed: trichoid, basiconic, coeloconic, styloconic and sensory pit. Only trichoid sensilla are found on the scape of the antenna, while both trichoid and styloconic sensilla are located on the antennal pedicels of all species studied. Basiconic sensilla are the most numerous of the sensilla found on the antennae of both sexes of all fly species studied and are comprised of two subtypes: large and small basiconic sensilla. Coeloconic sensilla are characterized by short pegs, with either grooved or smooth surfaces, that are sunken into deep depressions. No marked difference was observed in the number, morphological structure or distributional pattern of any of the sensilla among the species studied, with the exception of there being more numerous sensory pits detected in female P. dux compared to the other species. The suggested function of each antennal sensillum was based on comparison with results of other investigations on similar sensilla.     

Antennal sensillae of five stored-product psocids pests (Psocoptera: Liposcelididae)

Psocids are important pests of stored-products. In this study, the external morphology, types, and distribution of antennal sensillae in both male and female adults of five psocid species, Liposcelis bostrychophila, L. entomophila, L. tricolor, L. decolor, and L. paeta (Psocoptera: Liposcelididae) using scanning electron microscopy. The filiform antennae of the liposcelids are made up of the scape, pedicel, and a long flagellum composed of thirteen flagellomeres. Four morphological types of sensillae are recorded in both male and female species, including Böhm bristles, chaetal sensillae, microtrichial sensillae, and basiconic sensillae. The abundance and distribution of four of these sensilla types on the antennae of male and female are not significantly different in the five psocids species studied. In addition, the possible functions of the above sensilla types are discussed in light of previously published literature; mechanoreception (Böhm bristles and microtrichial sensillae) and olfaction (chaetal and basiconic sensillae). Future functional antennal morphology and electrophysiological studies are needed to confirm these proposed functions.     

The antennal sensilla of adult mayflies: Rhithrogena semicolorata as a case study

The present paper reports on an ultrastructural investigation of the sensilla on the antennal flagellum of the imago (SEM and TEM) and subimago (SEM) of the mayfly Rhithrogena semicolorata. Sensilla coeloconica of two different sizes, named big and small pegs, are distinguishable in both stages. The big pegs are lobe-shaped with a porous cuticle. The small pegs are more or less conical with a smooth cuticle. TEM seriate sections revealed that the big pegs are innervated by three neurons, which enter the peg. The dendrite sheath opens at the entrance to the peg and leaves the dendrites in contact with the outside through pores and pore tubules. This structure is in agreement with that reported for single-walled olfactory receptors. The small pegs may be innervated by two or four neurons. In both cases, two unbranched dendrites enter the peg and are in close association with the peg wall. These show features typical of hygroreceptors. In the small pegs innervated by four neurons, the other two neurons stop short of entering the peg, and one of them branches. This last could constitute the thermoreceptive unit. The function of the fourth neuron is unknown.The presence of olfactory sensilla in adult mayflies, such as Paleoptera, is relevant to the study of evolutionary trends in insect odour perception. Hygro–thermoreceptors are important in such fragile aquatic insects, which need a moist environment for moulting and reproduction.     

Sensilla of the antenna and palp of Hydrotaea chalcogaster (Diptera: Muscidae)

Hydrotaea chalcogaster is a fly species of medical and forensic importance in many parts of the world. In this study, we investigated the sensilla of the antenna and palp of the adult female fly using scanning electron microscopy. The antennal scape has one type of sensillum, the sharp-tipped sensillum trichodeum; whereas, the antennal pedicel also possessed this type of sensillum in addition to an unidentified type. Three types of sensilla were found on the flagellum: (1) sensilla basiconica, with both large and small sensilla basiconica showing wall pores, (2) sensilla coeloconica, with a smooth surface, and (3) sensory pits, with wall pores of pegs. The arista is located dorso-laterally on the flagellum and has three segments. Short microtrichia are located around the distal end of its second segment and on the proximal half of the third segment. Both large sharp-tipped sensilla chaetica and small sensilla basiconica with wall pores were observed on the palps. Results of this study contribute to our overall understanding of the ultrastructural morphology of sensilla on the antenna and palp of H. chalcogaster.     

Antennal sensilla of the decapitating phorid fly, Pseudacteon tricuspis (Diptera: Phoridae)

Pseudacteon tricuspis Borgmeier is a parasitic decapitating phorid fly (Diptera: Phoridae), which has been released in the southern United States in the last decade for biological control of invasive imported fire ants, Solenopsis spp. In a previous study, we demonstrated that P. tricuspis uses fire ant semiochemicals to locate host worker ants. To provide a solid background in support of our research on mechanisms of host location and olfaction in phorid flies, we studied the morphology of the antennal sensilla of both sexes of P. tricuspis using scanning and transmission electron microscopy. Antennae of P. tricuspis show strong sexual dimorphism in structure and shape: the female has a feathered arista which is located distally on the flagellum and has three sub-segments that bear small spinules of microtrichia. The first two antennal segments, scape and pedicel, bear no sensillum, but were densely covered by microtrichia. Three major types of sensilla were found on the flagellum (funicle) of both sexes: sensilla trichodea, sensilla basiconica, and sensilla coeloconica. Two of these, s. trichodea and s. basiconica were differentiated into three (short, medium, and long) and two (blunt-tip and sharp-tip) subtypes, respectively, for a total of six morphologically different sensilla subtypes. Ultrastructural studies revealed multiple wall pores on s. trichodea and s. basiconica, possibly suggesting chemoreceptory functions. The third and least abundant sensilla type, s. coeloconica, had no wall pores and may function as thermo-hygroreceptors. With the exception of the long subtype of s. trichodea which was recorded only on the male antennae, no other marked sexual differences were recorded in the number and distribution of antennal sensilla in P. tricuspis. These results are discussed in relation to the host location behavior of P. tricuspis, and could facilitate future studies on the neurobiology of olfaction in decapitating phorid flies.     

Estimation of the specific mass effect in the isotope shifts of energy levels in the optical spectrum of Ba I and Ba II

A graphical method for separating mass and volume effects from purely optical isotope shift data is presented and compared with King's “bunching” method. Recent experimental data on isotope shifts for a wide range of spectral lines in the naturally abundant isotopes of Ba I and Ba II are analysed. Some agreement is found with muonicx-ray data concerning the nuclear size for the isotopes 136, 137 and 138, but there is disagreement (over 20%) for the other isotopes. Thelevel isotope shifts are further parameterised in terms of a linear model, and the specific mass effect is decomposed into sums of one-electron and two-electron shift parameters with respect to the inert-gas like ground state of Ba III.     

The epipharyngeal sensilla of the damselfly Ischnura elegans (Odonata,Coenagrionidae)

The knowledge on Odonata adult mouthparts sensilla is scanty and, notwithstanding the epipharynx in the labrum is considered an organ of taste, no ultrastructural investigation has been performed so far on this structure in Odonata. The labrum of the adult of the damselfly Ischnura elegans (Odonata, Coenagrionidae) shows on its ventral side the epipharynx with sensilla represented by articulated hairs and by small pegs located at the apex of slightly raised domes. Under scanning and transmission electron microscope, the articulated hairs, with a well developed socket and tubular body, have the typical structure of bristles, the most common type of insect mechanoreceptors, usually responding to direct touch; the pegs, showing an apical pore together with a variable number of sensory neurons (from two to five), the outer dendritic segments of which show a dendrite sheath stopping along their length, have features typical of contact chemoreceptors.