The antennae of damselfly larvae

The larval antennal sensilla of two Zygoptera species, Calopteryx haemorroidalis (Calopterygidae) and Ischnura elegans (Coenagrionidae) are investigated with SEM and TEM. These two species have different antennae (geniculate, setaceous) and live in different environments (lotic, lentic waters). Notwithstanding this, similarities in the kind and distribution of sensilla are outlined: in both species the majority of sensilla types is located on the apical portion of the antenna, namely a composed coeloconic sensillum (possible chemoreceptor), two other coeloconic sensilla (possible thermo-hygroreceptors) and an apical seta (direct contact mechanoreceptor). Other mechanoreceptors, such as filiform hairs sensitive to movements of the surrounding medium or bristles positioned to sense the movements of the flagellar segments, are present on the antenna. Similarities in the antennal sensilla types and distribution are observed also with other dragonfly species, such as Onychogomphus forcipatus and Libellula depressa. A peculiar structure with an internal organization similar to that of a gland is observed in the apical antenna of C. haemorroidalis and I. elegans and it is present also in O. forcipatus and L. depressa. The possible function of this structure is at the moment unknown but deserves further investigations owing to its widespread presence in Odonata larvae.     

The antennal lobe of Libellula depressa (Odonata,Libellulidae)

Here we describe the antennal lobe of Libellula depressa (Odonata, Libellulidae), identified on the basis of the projections of the afferent sensory neurons stemming from the antennal flagellum sensilla. Immunohistochemical neuropil staining as well as antennal backfills revealed sensory neuron terminal arborizations covering a large portion of the antennal lobe. No clear glomerular structure was identified, thus suggesting an aglomerular antennal lobe condition as previously reported in Palaeoptera. The terminal arbors of backfilled sensory neurons do, however, form spherical knots, probably representing the connections between the few afferent neurons and the antennal lobe interneurons. The reconstruction revealed that the proximal part of the antennal nerve is divided into two branches that innervate two spatially separated areas of the antennal lobe, an anterioventral lobe and a larger posteriodorsal lobe. Our data are consistent with the hypothesis that one tract of the antennal nerve of L. depressa contains olfactory sensory neurons projecting into one of the sublobes, while the other tract contains thermo-hygroreceptive neurons projecting into the other sublobe.     

Olfaction in dragonflies: electrophysiological evidence

The problem of olfaction in Paleoptera (Odonata, Ephemeroptera) cannot be considered fully elucidated until now. These insects have been traditionally considered anosmic, because their brain lacks glomerular antennal lobes, typically involved in Neoptera odor perception. In order to understand if the presumed coeloconic olfactory receptors described on the antennal flagellum of adult Odonata are really functioning, we performed an electrophysiological investigation with electroantennogram (EAG) and single cell recordings (SCR), using Libellula depressa L. (Odonata, Libellulidae) as a model species. Odors representing different chemical classes such as (Z)-3-hexenyl acetate (acetate ester), (E)-2-hexenal, octanal (aldehydes), (Z)-3-hexen-1-ol (alcohol), propionic acid, butyric acid (carboxylic acids), and 1,4-diaminobutane (amine) were tested. Most of the tested chemicals elicited depolarizing EAG responses in both male and female antennae; SCR show unambiguously for the first time the presence of olfactory neurons in the antennae of L. depressa and strongly support the olfactory function of the coeloconic sensilla located on the antennal flagellum of this species. Electrophysiological activity may not necessarily indicate behavioral activity, and the biological role of olfactory responses in Odonata must be determined in behavioral bioassays. This study represents a starting point for further behavioral, electrophysiological, neuroanatomical and molecular investigation on Odonata olfaction, a research field particularly interesting owing to the basal position of Paleoptera, also for tracing evolutionary trends in insect olfaction.     

Ultrastructure of the antennal sensilla of the mayfly Baetis rhodani (Pictet) (Ephemeroptera : Baetidae)

The distribution and fine morphology of antennal sensilla of nymphal and adult mayfly, Baetis rhodani (Ephemeroptera : Baetidae), were examined. In the nymph, various kinds of sensilla (chaetica, basiconica, coeloconica and cuticular pits) are differently arranged on the antennal segments, whereas sensilla campaniformia delimit the distal border of the pedicel. A peculiar kind of sensillum basiconicum, named flat-tipped sensillum, is present along the entire antenna, even though in the flagellum it has a regular arrangement between the cuticular lobes that delimit the distal border of each article. In the subimago the scape and pedicel are profusely covered with microtrichia and scattered sensilla trichodea, whereas the flagellum shows cuticular ribs. Sensilla coeloconica are present along the ventral side of the flagellum. In the imago, the antenna is completely decorated with scales among which sensilla trichodea and sensilla coeloconica occasionally occur. As in the nymph, adult mayflies have a ring of sensilla campaniformia along the distal border of the pedicel. When compared with nymphal antennae, those of adults have fewer types of sensilla, presumably in relation to the short, non-feeding terrestrial life.     

Antennal sensilla of whiteflies: Trialeurodes vaporariorum (Westwood), the glasshouse whitefly,and Aleyrodes proletella (Linnaeus), the cabbage whitefly, (Homoptera : Aleyrodidae). Part 2: Ultrastructure

A transmission electron microscope study of the antennal sensilla of the whitefly Trialeurodes vaporariorum and Aleyrodes proletella (Homoptera : Aleyrodidae) revealed that of the sensilla unique to the antennal flagellum (basiconic, coeloconic and small digitate-tipped sensory pegs), basiconic and coeloconic sensilla occur as subtypes. Four subtypes of basiconic cone sensilla occur on the flagella of T. vaporariorum and 3 on A. proletella. All the subtypes of basiconic sensilla have an ultrastructure typical of olfactory sensilla and probably have a primary olfactory function. Two subtypes of coeloconic sensilla occur on the flagella of both species. Their ultrastructure suggests primarily a chemosensory function. The digitate-tipped sensory peg of both species possesses a triad of neurones which have ultrastructural characteristics similar to the known thermo-/hygroreceptors of other insect species. The other sensilla, which occur on the antennae of the whiteflies, include cheatae, campaniform and subcuticular sensilla, all of which have an ultrastructure typical of mechanoreceptors.     

The sense of smell in Odonata: An electrophysiological screening

Volatile chemicals mediate a great range of intra- and interspecific signalling and information in insects. Olfaction has been widely investigated mostly in Neoptera while the knowledge of this sense in most basal insects such as Paleoptera (Odonata and Ephemeroptera) is still poor. In the present study we show the results of an electrophysiological screening on two model species, Libellula depressa (Libellulidae) and Ischnura elegans (Coenagrionidae), representatives of the two Odonata suborders Anisoptera and Zygoptera, with the aim to deep the knowledge on the sense of smell of this insect order. The antennal olfactory sensory neurons (OSNs) of these two species responded to the same 22 compounds (out of 48 chemicals belonging to different functional groups) encompassing mostly amines, carboxylic acids or aldehydes and belonging to green leaf volatiles, vertebrate related volatiles and volatiles emitted by standing waters bacteria. The properties of Odonata OSNs are very similar to those of ionotropic receptors (IRs) expressing OSNs in other insects.     

The antennal sensilla of the adult of Libellula depressa (Odonata: Libellulidae)

An ultrastructural investigation (SEM, TEM) on the antennal flagellum of the adult of the dragonfly Libellula depressa (Odonata:Libellulidae) revealed sensilla located in pits on the lateral-ventral side of the antenna. These sensilla are represented by sensilla coeloconica and by deeply sunken sensilla. The sensilla coeloconica are innervated by three unbranched dendrites, which enter the peg and show a dendrite sheath ending at the base of the peg. The peg has no socket and its cuticle is irregular with wide pore-like structures at the base of which actual pores are visible. The structure of these coeloconic sensilla is in agreement with that reported for single-walled insect chemoreceptors. The deeply sunken sensilla are represented by two kinds of sensilla styloconica, named type-1 and type-2, located at the bottom of deep cavities appearing as simple openings on the antennal surface. These sensilla are no-pore sensilla with inflexible socket and unbranched dendrites and, notwithstanding their structural differences, share common features typical of thermo/hygroreceptors. The presence of chemoreceptors in adult dragonflies sheds light on evolutionary trends in insect perception; the previously unknown occurrence of thermo/hygroreceptors in dragonflies is very important in view of the reported ability of Odonata to thermoregulate heliothermically.     

Antennal and ovipositor sensilla of Pseudoligosita yasumatsui (Hymenoptera: Trichogrammatidae)

Pseudoligosita yasumatsui Viggiani and Subba Rao 1978 (Hymenoptera: Trichogrammatidae) is a common egg parasitoid of rice insect pests. The surface morphology of the antenna and ovipositor on P. yasumatsui was examined using scanning electron microscopy. The antenna of P. yasumatsui is geniculate in shape, hinged at the scape-pedicel joint, approximately 190 μm in length and consists of seven antennomeres. In total, the male and female antennae have ten different types of sensilla: trichoid sensilla type 1, 2, 3, 4, 5, 6, campaniform sensilla, basiconic sensilla, and placoid sensilla type 1 and 2. The flagellum of the female antenna is covered with cuticular pores, which are absent on the male antennal flagellum. The distal extremity of its ovipositor stylet has campaniform sensilla and styloconic sensilla. Trichoid sensilla found on its apical abdomen part may play a role in the host detection and egg placement. The types and distribution of antennal and ovipositor sensilla on the parasitoid were discussed.     

Functional anatomy of sensory structures on the antennae of Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae)

The ultrastructure and distribution of sensilla on the antennae of the cabbage stem flea beetle, Psylliodes chrysocephala, were investigated using scanning and transmission electron microscopy techniques. Eight different sensillar types were distinguished. These were; hair plate sensilla, sensilla chaetica, three types of sensilla trichodea, sensilla basiconica, grooved peg sensilla and styloconic sensilla. The sensilla chaetica are known to be gustatory receptors. Ultrastructure indicates that the hair plate sensilla and sensilla trichodea type one are probably mechanoreceptors, whilst the sensilla styloconica are probably thermo-hygro receptors. These thermo-hygroreceptors are unusual in that they are innervated by two sensory cells (one hygroreceptor and one thermoreceptor) rather than the more usual triad. The remaining four sensillar types all have a porous hair shaft, indicating an olfactory role. One of these (the grooved peg sensillum) may also have a thermoreceptive function. No sexual dimorphism was found in the structure, number or distribution of the antennal sensilla.     

六斑月瓢虫触角及感觉器扫描电镜观察

[目的] 明确六斑月瓢虫雌雄成虫触角感觉器种类、分布及形态特征。[方法] 利用扫描电子显微镜对六斑月瓢虫雌、雄成虫触角形态及触角感受器超微结构进行观察。[结果] 六斑月瓢虫成虫触角由柄节、梗节和鞭节组成,柄节长度与宽度显著大于梗节长度与宽度;鞭节分为9个亚节,末端3节横向膨大呈锤状。雌雄成虫触角上共有8种感觉器:刺形感觉器(SC)、毛形感觉器(ST)、锥形感觉器(SB)、腔形感觉器(CaS)、钟形感觉器(CS)、哑铃形感觉器(DS)、香肠形感觉器(SS)及B?hm氏鬃毛感觉器(BB)。以毛形感觉器和刺形感觉器分布最广,遍布触角;B?hm氏鬃毛仅存在于触角柄节与梗节;触角鞭节第9亚节顶端密布7种触角感觉器。六斑月瓢虫雌雄成虫触角长度、触角感觉器类型及分布无显著差异。[结论] 六斑月瓢虫成虫触角上共有8种感觉器,其触角可能具有感知机械刺激、识别化学信息素及感受温湿度变化的作用。本研究为进一步了解六斑月瓢虫触角与其行为间的关系提供基础资料。     

Morphological and functional heterogeneity in olfactory perception between antennae and maxillary palps in the pumpkin fruit fly,Bactrocera depressa

The morphology and ultrastructure of the olfactory sensilla on the antennae and maxillary palps were investigated through scanning electron microscopy (SEM) and transmission electron microscopy (TEM), and their responses to five volatile compounds were measured using electroantenogram (EAG) and electropalpogram (EPG) techniques in the pumpkin fruit fly, Bactrocera depressa (Shiraki; Diptera: Tephritidae). Male and female B. depressa displayed distinct morphological types of olfactory sensilla in the antennae and maxillary palps, with predominant populations of trichoid, basiconic, and coeloconic sensilla. Basiconic sensilla, the most abundant type of olfactory sensilla in the antennae, could be further classified into two different types. In contrast, the maxillary palps exhibited predominant populations of a single type of curved basiconic sensilla. High‐resolution SEM observation revealed the presence of multiple nanoscale wall‐pores on the cuticular surface of trichoid and basiconic sensilla, indicating that their primary function is olfactory. In contrast, coeloconic sensilla displayed several longitudinal grooves around the sensillum peg. The TEM observation of individual antennal olfactory sensilla indicates that the basiconic sensilla are thin‐walled, while the trichoid sensilla are thick‐walled. The profile of EAG responses of male B. depressa was different from their EPG response profile, indicating that the olfactory function of maxillary palps is different from that of antennae in this species. The structural and functional variation in the olfactory sensilla between antennae and maxillary palps suggests that each plays an independent role in the perception of olfactory signals in B. depressa.     

Sense organs on the antennal flagellum of the green stink bug,Nezara viridula (L.) (Heteroptera : Pentatomidae): Sensillum types and numerical growth during the post-embryonic development

The sensilla on the antennal flagellum of Nezara viridula (Heteroptera : Pentatomidae) were studied with the scanning and transmission electron microscopes. Seven morphologically distinct types of sensilla were identified in adults. The number and distribution of six types were studied. Male and female antennae bear the same numbers and types of sensilla. As in other Heteroptera, the flagellum comprises two segments. The second flagellar segment bears more sensilla than the first segment. Sensilla are evenly distributed over the flagellum surface. The total number of sensilla increases during the post-embryonic development. Individual numerical growths of each type follow simple exponential laws when expressed as functions of developmental stage. The growth in number shows positive allometry in respect to body size for sensillum types 1, 2 + 5 and 4, but negative allometry for types 3 and 6. The density of sensilla remains constant. These findings are compared with previously published data from other heterometabolous insects. Adaptation to the increase in body size during the post-embryonic development seems to be the main determinant of the growth of the olfactory system in N. viridula.     

The chemosensory system of the monophagus grasshopper,Bootettix argentatus bruner (Orthoptera : Acrididae)

Bootettix argentatus (Orthoptera : Acrididae) is a monophagous grasshopper in the subfamily Gomphocerinae. The numbers of chemoreceptors in most groups on the mouthparts increase with each molt and are generally similar in the 2 sexes. There are fewer sensilla in the A3 groups than in other gomphocerine grasshoppers, and the possibility that this is related to host-plant specificity is discussed. Only very small numbers of sensilla are present on the pulvillar pads of the tarsi and on the arolia. The number of sensilla on the antennae is proportional to the length of the flagellum and coeloconic sensilla comprise 30–35%. Most of the remainder are multiporous basiconic sensilla. Adult males have more antennal sensilla than adult females.     

椰心叶甲啮小蜂复眼和触角在交配中的作用及其超微结构的扫描电镜观察

对寄生蜂交配行为的了解将有助于发展对其行为调控的技术,提高寄生蜂对害虫控制的效能。为探讨椰心叶甲啮小蜂Tetrastichus brontispae Ferrière的复眼和触角在交配中的作用,用水溶性黑色素和液体石蜡分别涂抹该蜂复眼和触角后观察其交配行为,并利用扫描电镜（SEM）观察其复眼和触角的超微结构,分析雌、雄蜂触角感器的分布与数量差异。结果表明：椰心叶甲啮小蜂雄蜂的复眼在交配过程中起重要作用,雌蜂复眼作用不显著。在椰心叶甲啮小蜂求偶识别和接受过程中,雄蜂触角柄节部位起主要作用,其次是棒节部位,再次是鞭节的索亚节部位,而雌蜂触角鞭节索亚节部位起主要作用,然后是棒节部位,最后是柄节部位。扫描电镜观察表明椰心叶甲啮小蜂触角上共有8种感器,其中毛形感器和板形感器是主要感器,雌、雄蜂触角有明显的性二型现象,表现为触角大小不同及触角感器类型、大小、数量、分布不同。     

Morphology of antennal sensilla of the brown spruce longhorn beetle,Tetropium fuscum (Fabr.) (Coleoptera: Cerambycidae)

The antennal sensilla of the brown spruce longhorn beetle, Tetropium fuscum (Fabr.) (Coleoptera: Cerambycidae) were examined with particular focus on the sensilla present on the apical flagellomere. T. fuscum antennae are composed of 11 segments, namely the scape, pedicel, and nine flagellomeres. Nine types of sensilla were observed: three types of sensilla chaetica, sensilla trichodea, two types of sensilla basiconica, grooved peg sensilla, thick-walled sensilla, and Böhm bristles. Seven of these types were present on the apical flagellomere, the exceptions were sensilla chaetica type 3 and Böhm bristles. There were no significant differences in the distribution or density of sensilla present on the ninth flagellomere of males and females, except that males had significantly more sensilla chaetica type 1, which are put forward as the putative contact chemoreceptors for T. fuscum.     

Comparison of the antennal sensilla of females of four fig-wasps associated with Ficus auriculata

A comparison was performed of the antennal sensilla of females of four chalcid wasp species Ceratosolen emarginatus Mayr, 1906, Sycophaga sp., Philotrypesis longicaudata Mayr, 1906, and Sycoscapter roxburghi Joseph, 1957, which are specific and obligatory associated with Ficus auriculata (Lour, 1790). The four species exhibit different oviposition strategies in the fig ovules where their offspring hatch and develop. Antennal sensilla morphology was evaluated using scanning electron microscopy. Females of the four species present 11 morphologically similar types of sensilla: trichoid sensilla, sensilla obscura, chaetica sensilla 1 and 2, which all have mechanosensory functions; uniporous basiconic sensilla, which are presumably contact chemosensilla; basiconic capitate peg sensilla, coeloconic sensilla 1, multiporous basiconic and placoid sensilla, which may be regarded as olfactory sensilla, and coeloconic sensilla 2 and 3, which are presumed to be proprioreceptors or pressure receptors. The four species have significant differences in the abundance and arrangement of trichoid sensilla and chaetica sensilla 1 on the flagellum. The coeloconic sensilla and sensilla obscura only occur on the antennae of C. emarginatus that enter figs. The chemosensilla which are presumably involved in host discrimination, i.e., basiconic sensilla, multiporous placoid sensilla and basiconic capitate peg sensilla, are similar in shape and configuration, although they present some differences in abundance. These findings provide practical information on the adaptations of fig wasps and the relationship between multisensory antennae and functions in fig wasp behaviour.     

桉小卷蛾触角感器的超微结构

[目的] 桉小卷蛾是桉树、番石榴等林木和果树的一种重要害虫。为确定桉小卷蛾成虫触角上的感器对寻找生存环境及对外界环境的各种刺激所起的作用,研究了桉小卷蛾触角感器的种类、形态和分布。[方法] 利用扫描电镜对桉小卷蛾成虫触角感器进行了详细观察和研究。[结果] 桉小卷蛾雌、雄蛾触角由柄节、梗节和鞭节组成,触角的外侧面覆盖有鳞片;雄性触角略短于雌虫,且鞭节近1/4处有一凹陷。共发现7种感器,即鳞形感器、毛形感器、锥形感器、腔锥形感器、耳形感器、刺形感器和Böhm氏鬃毛,其中以毛形感器数量最多。[结论] 桉小卷蛾雌雄成虫触角的感器种类、分布、数量有一定的规律但又有差异。建议应用气相-触角电位联用技术GC-EAD和触角电位EGA等技术进一步解析触角感器的功能。     

Scanning Electron Microscopy Studies of the Antennal Sensilla of Metaphycus parasaissetiae Zhang & Huang (Hymenoptera: Encyrtidae)

Metaphycus parasaissetiae Zhang & Huang (Hymenoptera: Encyrtidae) is an important adult parasitoid of Parasaissetia nigra Nietner (Hemiptera: Coccoidea). The external morphology of the antennal sensilla of male and female M. parasaissetiae was examined using scanning electron microscopy. The geniculate antennae of male and female M. parasaissetiae were composed of a scape with a basal radicula, a barrel-shaped pedicel, and a long flagellum. Twelve morphologically distinct types of sensilla were identified, including multiporous placoid sensilla, campaniform sensilla, finger-like sensilla, multiporous basiconic sensilla (BS-1), three aporous types of basiconic sensilla (BS-2, BS-3, and BS-4), two types of aporous trichoid sensilla (TS-1 and TS-3), a type of multiporous trichoid sensilla (TS-2), and two types of sensilla chaetica (CH-1 and CH-2). Sex dimorphism in the sensilla composition of M. parasaissetiae is also observed. Major differences between the sexes were found in the number, distribution, shape, structure, and size of the identified sensilla. We also discuss on the functional aspects of these sensilla to elucidate the mechanisms involved in host searching and courtship behavior of M. parasaissetiae.     

External morphology of antennal sensilla of trichogramma australicum girault (Hymenoptera : Trichogrammatidae)

External morphology of antennal sensilla on female and male Trichogramma australicum (Hymenoptera : Trichogrammatidae) was examined using scanning electron microscopy. Antennae show strong sexual dimorphism in structure and types of sensilla. The female antenna displays 14 types of sensilla: basiconic capitate peg sensilla (types 1 and 2), campaniform sensilla, chaetica sensilla (types 1–3), coeloconic sensilla, falcate sensilla, placoid sensilla (types 1 and 2), styloconic sensilla and trichoid sensilla (types 1–3). The male antenna displays 12 types of sensilla: basiconic capitate peg sensilla (type 2), campaniform sensilla, chaetica sensilla (types 1–5), coeloconic sensilla, placoid sensilla (type 1), and trichoid sensilla (types 3–5). Falcate and styloconic sensilla occur only on the female antenna. Both sensilla probably are associated with host examination, host discrimination and oviposition behaviour. Male antennal trichoid sensilla types 4 and 5 are probably associated with courtship behaviour, because these types occur only on the male. We propose the term “falcate sensilla” for a unique female antennal sensilla; the number of falcate sensilla may be used for identification of Trichogramma spp. In addition, we report the presence of placoid sensilla type 2 and difference in structure of coeloconic sensilla in T. australicum. Variation in structure and position of antennal sensilla are discussed.     

Antenna morphology and sensilla microstructure of the male and female scarab beetle,Lepidiota mansueta Burmeister (Coleoptera: Scarabaeidae)

Studies on Lepidiota mansueta Burmeister were conducted to describe and compare the external morphology of the antenna and its sensilla of male and females. Distinct variations were observed in antennal segments between the sexes with males possessing longer scape, pedicel, flagellum and lamellar club segments. Variation was also seen in the antennal sensilla, with males possessing three different types of sensilla while the females possessed seven types of sensilla. Sensilla trichodea (TRS), sensilla placodea (PLAS) type I and sensilla placodea (PLAS) type II were common in both sexes while sensilla chaetica (CHS), sensilla auricilica (AUS) type I, sensilla auricilica (AUS) type II and sensilla auricilica (AUS) type III were found only in females.