The antennae of neopseustid moths: Morphology and phylogenetic implications, with special reference to the sensilla (Insecta, Lepidoptera, Neopseustidae)

The antennae of Lepidoptera Neopseustidae were examined with the scanning electron microscope. The studied species, Nematocentropus cfr. omeiensis, Neopseustis meyricki, Synempora andesae, Apoplania valdiviana and Apoplania penai possess nine types of antennal flagellum sensilla: multiporous large sensilla basiconica, multiporous thin sensilla basiconica, multiporous small sensilla basiconica, multiporous sensilla trichodea, multiporous sensilla coeloconica; uniporous sensilla chaetica; aporous sensilla chaetica, aporous stylus-shaped sensilla chaetica, aporous sensilla styloconica.The multiporous sensillum type here termed “multiporous large sensillum basiconicum” is unknown from other Lepidoptera and probably constitutes an autapomorphy of the family Neopseustidae. This sensillum type is remarkable by having a single base in female Apoplania and Synempora while in male Apoplania it has a bifid or trifid base, and in male Synempora it is composed of two or three incompletely separated hairs. This may be the first recorded example of a sexually dimorphic lepidopteran sensillum type. The stylus-shaped sensillum chaeticum is a primitive type which occurs only in some lower Lepidoptera.     

Morphology and distribution of sensilia on the cephalic appendages,tarsi and ovipositor of the European sunflower moth,Homoeosoma nebulella Den. & Schiff. (Lepidoptera : Pyralidae)

Sensilla on male and female antennae, mouthparts, tarsi and on the ovipositor of the European sunflower moth, Homoeosoma nebulella (Lepidoptera : Pyralidae) have been investigated by means of scanning electron microscope. The antennal flagellum bears multiporous sensilla (trichodea, basiconica, auricillica, coeloconica) and uniporous sensilla (chaetica, styloconica). A sexual dimorphism is found in the numbers of sensilla trichodea (8300 in males and 6000 in females) and sensilla basiconica (1150 in females and 650 in males). The proboscis has uniporous s. styloconica, multiporous s. basiconica (described for the first time in Lepidoptera), aporous s. chaetica, and some rare uniporous s. basiconica in the food channel. The labial palp-pit organ houses multiporous s. coeloconica, while the maxillary palps have aporous sensilla. Tarsi of both sexes possess one type of aporous s. chaetica and 2 types of uniporous s. chaetica. There is no significant difference in the distribution of tarsal sensilla between males and females. The surface of each lobe of the ovipositor is covered by 3 types of aporous s. trichodea of different lengths and about 60 multiporous s. basiconica, sensilla unusual on the ovipositor of Lepidoptera. The role of sensilla in the oviposition site selection is discussed.     

红火蚁触角及其上感受器的扫描电镜观察

应用扫描电镜对采自我国广东吴川和广西南宁的红火蚁Solenopsis invicta Buren工蚁、有翅雌蚁和雄蚁触角的形态和感受器类型进行了研究。结果表明,工蚁、雌蚁和雄蚁的触角存在着较大的差异。工蚁和雌蚁的触角为膝状,末端2或3节膨大为棒状,雌蚁的触角11节,比工蚁(10节)的多1节。雄蚁的触角为线状,12节,末端没有棒状结构。工蚁和雌蚁的触角上共有7种感受器,分别为毛形、曲毛形、锥形、腔锥形、坛形、刺形感受器和Böhm氏鬃毛,前6种感受器大多集中在触角末端的棒节,棒节以外的各鞭节上主要是毛形感受器,Böhm氏鬃毛则分布于柄节和梗节的基部,鞭节上无此感受器。雄蚁触角上也有7种感受器,但其分布与工蚁的有较大差异,Böhm氏鬃毛的分布与工蚁和雌蚁的相同,锥形感受器分布在末端2节,坛形感受器分布在末端3节,其余的4种感受器较为均匀地分布在各鞭节,而不是聚集在端节。另外,新发现雄蚁中存在一种与已知的毛型感受器不同的超短毛状感受器,定名为超短毛感受器。最后,对红火蚁的触角及感受器在蚁群社会性行为中的作用进行了讨论。     

Perception of potential sex pheromones and host-associated volatiles in the cotton plant bug,Adelphocoris fasciaticollis (Hemiptera: Miridae): morphology and electrophysiology

Highly developed chemoreception allows insects to detect foods, find mates, and escape natural enemies. We described the structures and distributions of antennal chemosensilla in Adelphocoris fasciaticollis Reuter by scanning electron microscopy and transmission electron microscopy. Seven major types of antennal sensilla were identified in adults of both sexes. Types 1 and 2 are sensilla chaetica and have thick cuticular walls with conspicuous grooves at their surfaces. Types 3 and 4 are multiporous sensilla trichodea with 1–3 dendrites located at the sensillum lymph, indicating a putative olfactory function. Types 5 and 6 are typical sensilla basiconica but share different characteristics in both external morphology and internal ultrastructure, and may be involved in the perception of host-associated odorants. The last sensilla were Böhm bristles. In addition to the morphological characterization, electrophysiological responses of antennal chemosensilla to 51 semiochemicals were investigated based on electroantennogram (EAG) recordings. Results revealed that different chemical stimuli elicited significantly different dose-dependent EAG responses, in which potential sex pheromone components and green leaf volatiles showed relatively higher EAG responses, but neither monoterpenes nor sesquiterpenes can elicit favorable EAG values. The results provided direct morphological and electrophysiological evidence that the adult antennae of A. fasciaticollis could function in searching for mates and host plants.     

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.     

Mate recognition and antennal morphology of Octodonta nipae (Coleoptera: Chrysomelidae) adults

The nipa palm hispid beetle, Octodonta nipae (Maulik) has been killing palm trees since its introduction into Hainan province, China, from Malaysia in 2001. It continues to spread within Hainan province, northeast to Fujian province, and northwest to Yunnan province within China. Knowledge on signals involved in mate location and recognition could help develop effective integrated pest management programs. In the present study, we first experimentally proved that antennae were essential in success of O. nipae mating. We then excised various segments/flagellomeres of adult male and female antennae and observed their mating behavior. Results revealed that the 5th to 9th flagellomeres, especially those of males, were important for the mating success. Finally, in an attempt to elucidate the types of antennal sensilla accountable for the mating success, morphology of O. nipae antennae was studied in detail with scanning electron microscopy. Six types of sensilla were distinguished: aporous sensilla trichodea (T1), multiporous sensilla trichodea (T2), aporous sensilla chaetica (Ch1), uniporous sensilla chaetica (Ch2), multiporous sensilla basiconica (B), and Böhm sensilla (Bm). Aporous sensilla trichodea is the most abundant; multiporous sensilla trichodea and sensilla basiconica are considered as olfactory receptors, and uniporous sensilla chaetica as gustatory receptor. Importance of flagellomeres 5–9 in mating success seemed to correspond to the abundance of sensilla on these segments.     

椰心叶甲啮小蜂触角感觉器的扫描电镜观察

应用扫描电镜(SEM)对椰心叶甲啮小蜂Tetrastichus brontispae Ferrière成虫触角感器的超微结构进行观察。结果表明,在触角上共有8种感器,分别为毛形感器、多孔板状感器、刺形感器、锥形感器、耳形感器、乳状感器、腔形感器和Bhm氏鬃毛;其中毛形感器和板形感器是主要感器,数量较多分布较广;雌雄蜂触角有明显的性二型现象,表现为雌雄触角大小不同,触角感器类型、大小、数量、分布不同。     

Antennal sensilla of male and female of the nut weevil,Curculio nucum Linnaeus, 1758 (Coleoptera: Curculionidae)

The nut weevil, Curculio nucum (Linnaeus, 1758), is the main pest in hazelnut orchards (Corylus avellana L.). Semiochemicals are interesting bio control tools that could be used to manipulate the pest behaviour and to control pest populations. The study of the sensorial equipment of the insect antennae provides information on the importance of olfaction in the adult life for host plants and mate findings as well as on the putative other senses. Before electrophysiological investigation, the knowledge of antennae equipment is also necessary. The aim of this study is to determine the types, number and location of sensilla on the antennae of male and female adult C. nucum in order to determine their implication in seeking a sexual partner and a host plant. The 12-segmented antenna comprises a scape, a 7-segmented funicle and a 4-segmented club. Out of the nine sensillum types listed, three are present on the scape and the funicle and seven types on the club which gathers 71–73% of the total of sensilla. Tactile aporous sensilla chaetica C1, gustatory uniporous sensilla chaetica C2, olfactory multiporous sensilla basiconica B1 and B2 are found on both the dorsal and ventral surfaces of the club in both sexes. Thermo-hygroreceptive dome-shaped sensilla D, olfactory multiporous sensilla basiconica B3 and olfactory multiporous fluted sensilla basiconica F are found exclusively on the ventral surface of the club, suggesting that these sensilla are utilized in host plant acceptance during antennal tapping. The sexual dimorphism concerns only the numbers of sensilla chaetica C1 and sensilla basiconica B2.     

Scanning Electron Microscopy Study of the Antennal Sensilla of <Emphasis Type="Italic">Monema flavescens</Emphasis> Walker (Lepidoptera: Limacodidae)

Monema flavescens Walker (Lepidoptera: Limacodidae) is a serious polyphagous defoliator. Using scanning electron microscopy, the external morphology of the antennal sensilla of this pest was examined for a better understanding of the mechanisms of insect-insect and insect-plant chemical communications. The antennae of M. flavescens were filiform in shape, and 11 morphological types of sensilla were found in both sexes. Six types of likely chemosensory sensilla were identified: uniporous sensilla chaetica, multiporous sensilla trichodea, and four types of multiporous sensilla basiconica. The sensilla identified as likely mechanoreceptors included two subtypes of aporous sensilla chaetica, aporous sensilla coeloconica, aporous sensilla styloconica, and Böhm’s bristles, whereas the position of the antennae was monitored by Böhm’s bristles.     

Antennal sensilla in five Psychodini moth flies (Diptera: Psychodidae: Psychodinae) pollinators of Arum spp. (Araceae)

The pollination of the genus Arum (Araceae) is mainly achieved by deception, the floral odour mimicking the pollinator ovipositing site. In order to discover the sensory organs involved in this attraction, we have studied the antennae of five species of psychodine moth-flies (former Psychoda sensu lato=Psychodini), pollinators of Arum spp. The antennae of the five Psychodini reveal seven types of sensilla: multiporous tribranched sensilla basiconica (sensilla ascoidea), multiporous sensilla basiconica, multiporous sensilla coeloconica, multiporous sensilla auricillica, uniporous sensilla basiconica, aporous sensilla chaetica, aporous Böhm’s sensilla. Each species possesses three, five or six of these sensillum types. All the multiporous sensilla are probably olfactory receptors while the uniporous sensilla basiconica must possess a contact chemoreceptive function. The multiporous tribranched sensilla basiconica (s. ascoidea), present in all the species, are the best candidates for the reception of the odours given off by the ovipositing sites and the inflorescences of Arum. The multiporous sensilla basiconica and the multiporous sensilla coeloconica may be involved respectively as CO₂ receptors or thermoreceptors. Psychoda phalaenoides, which is the main pollinator of A. maculatum, is the species which possesses the largest number of antennal sensilla. The sexual dimorphism, studied only in Psycha grisescens, as concerns the number of sensilla and the absence of a sensillum type which differ according to sex, is difficult to interpret.     

伪鞘榕小蜂雌雄成虫触角感器的超微形态、分布及适生意义

伪鞘榕小蜂Sycoscapter trifemmensis是一种寄生于鸡嗉子榕间花期榕果的专性寄生蜂,雌雄两性繁殖策略分化明显,为更好理解和诠释雌蜂寄主定位和雄蜂配偶识别机制,有必要对两性的触角感器进行观察。运用环境扫描电镜观察,对比和探讨了伪鞘榕小蜂雌雄成虫的触角和触角感器类型、分布、数量及其生态适应性。结果表明:雌蜂触角鞭节由11鞭小节组成,总长817.82±33.23μm,分布有毛形感器、刺形感器(类型1)、锥形感器(类型1)、多孔板形感器(类型1)、栓锥形乳突状感器5类5种;雄蜂触角鞭节仅由6鞭小节组成,全长为雌蜂的1/3,且各节有明显的缩短和增粗特征,着生感器包括毛形感器、刺形感器(类型2和类型3)、锥形感器(类型1和类型2)、多孔板形感器(类型2)、腔锥形感器5类7种。雌蜂触角感器的数量与分布显著高于雄蜂,且同类型感器在雌蜂上具有明显的延伸、增粗、分支的特征,以板形感器和锥形感器最为突出。伪鞘榕小蜂雌雄成虫的触角及其感器有明显的性二型,特别是与化学信息识别相关的感器,反映了雌雄蜂在不同生态环境和繁殖压力下的形态分化、行为策略和生态适应。     

Morphology and distribution of antennal sensilla in the pine wood nematode insect vectors Monochamus alternatus Hope and Monochamus saltuarius Gebler (Coleoptera: Cerambycidae)

As insect vectors of pine wood nematodes, Monochamus alternatus Hope and Monochamus saltuarius Gebler have different levels of attractiveness to semiochemicals. Although active use of chemical communication in both species is well known, there was no report on the differences in morphology and distribution of antennal sensilla between the two Monochamus species. The present study was carried out to explore the morphology, distribution, and ultrastructure of antennal sensilla of the two species using scanning and transmission electron microscopy. The overall shapes of the antennae were not different between M. alternatus and M. saltuarius. However, both M. alternatus and M. saltuarius exhibited sexual dimorphism overall. The antennae of each species were composed of the scape, pedicel, and nine flagellomeres, with four morphological types of sensilla: trichodea, basiconica, chaetica, and coeloconica. The four types occurred in varying numbers and distributions. The sensilla chaetica type was the most abundant in both species. Three types of sensilla (trichodea, basiconica, and chaetica) could be subdivided according to the two Monochamus species; sensilla trichodea could be divided into two subtypes, and sensilla basiconica and sensilla chaetica into three. All sensilla basiconica subtypes were multiporous, indicating that their primary function was olfactory. Sensilla trichodea subtype 2 was uniporous, indicating that the function of these sensilla might be gustatory. The morphological information obtained in our study provides a basis for future investigation of the sensory physiological function of each type of sensilla in the two species.     

Gall midge olfaction: Pheromone sensitive olfactory neurons in Contarinia nasturtii and Mayetiola destructor

This study describes the morphology and function of the antennal sensilla in two gall midge species, Contarinia nasturtii and Mayetiola destructor, where multi-component sex pheromones have been identified. Both species possess sensilla trichodea, s. coeloconica, s. chaetica and s. circumfila. Sensilla circumfila, which consist of several sensilla that bifurcate and fuse into one structure, are unique for the gall midges. In C. nasturtii s. circumfila are sexually dimorphic. In males, they form elongated loops suspended on cuticular spines, whereas in females they run like worm-like structures directly on the antennal surface. Single sensillum recordings demonstrated that olfactory sensory neurons housed in male s. circumfila in C. nasturtii responded to the female sex pheromone. In M. destructor, s. circumfila were attached to the antennal surface in both sexes, and displayed no response to sex pheromone components.A sexual dimorphism was also found in the number of s. trichodea per antennal segment in both C. nasturtii (male 1 vs. female 7) and M. destructor (male 13 vs. female 10). OSNs located in male M. destructor s. trichodea responded to the sex pheromone. This is the first gall midge single sensillum study, and the first demonstration of the functional significance of s. circumfila.     

中红侧沟茧蜂触角感受器的扫描电镜观察

利用扫描电镜对中红侧沟茧蜂Microplitis mediator的触角感受器进行了观察,发现了6个类型的感器,分别为毛形感器、板形感器、刺形感器、钟形感器、锥形感器、腔锥形感器。其中,毛形感器具有2种形态,锥形感器具有4种形态。钟形感器仅分布于雌蜂的触角上,锥形感器Ⅲ和Ⅳ仅分布于雄蜂的触角上。结合感受器的形态、分布和已报道的触角电位反应数据,对各感受器的功能进行了推测。     

黄脊竹蝗触角感器的电镜扫描观察

黄脊竹蝗是中国南方地区重要的竹子害虫,为了更好地理解黄脊竹蝗"趋尿行为"的生理生化机制,通过电镜扫描技术研究了黄脊竹蝗成虫触角感器的类型、数量、形态及分布特征,比较分析了其在雌、雄成虫间的差异。结果表明:黄脊竹蝗雌、雄成虫触角丝状,由1节柄节、1节梗节和23节鞭节构成,雌雄间触角长度及直径差异不显著(P>0.05);雌、雄成虫触角感器均有毛形感器Ⅰ、毛形感器Ⅱ、刺形感器、锥形感器Ⅰ、锥形感器Ⅱ和腔锥形感器6种;其中,锥形感器数量最多,约占感器总数的53%,主要分布在鞭节的第8～21亚节上;各类感器在雌、雄成虫间触角上的分布特征相似,雄成虫触角上感器总数、锥形和腔锥形感器数量显著多于雌成虫(P<0.05)。     

Morphology and distribution of antennal sensilla of the tarnished plant bug,Lygus lineolaris (Palisot de beauvois) (Hemiptera: Miridae)

Sensilla on the antennae of adult and last-instar nymphs of the tarnished plant bug, Lygus lineolaris (Hemiptera: Miridae), were examined with light, scanning and transmission electron microscopy. Six different types were identified in adult females and males and 5 types in last-instar nymphs: types 1 and 4 are sensilla trichodea, 2 and 3 are sensilla chaetica, and 5 and 6 are sensilla basiconica. Type 1 are located at distal region of terminal segment and type 2 are located at distal regions of proximal 3 segments in both adults and nymphs. Type 3 is present on all segments, more numerous on scape and pedicel and less abundant on distal third and fourth segments in both adult and nymphal stages. Types 4 and 6 are absent on the scape and present on the distal 3 antennal segments in adults, but they are present only on the distal-most antennal segment in nymphs. Type 5 sensilla are present only on second antennal segments in adults and are absent in nymphs. Sexual dimorphism is observed in total numbers: there are significantly more type(s) 3, 4, 5 and 6 sensilla in adult males than adult females. Types 1, 4 and 5 are multiporous with thin cuticle, branched dendrites and pore tubules which suggests an olfactory function. These sensilla have 3, 3 and 2 neurons, respectively. The type 6 sensillum has an apical pore and pores in the cuticular wall, and is innervated by 5 nerve cells with unbranched dendrites. Sensillar types 2 and 3 have thick cuticle, a single apical pore and nerve cells with unbranched dendrites. Type 2 has 1 neuron and type 3 has 2 chambers and 2 nerve cells.     

Fine structure and distribution of antennal sensilla of the desert locust, Schistocerca gregaria (Orthoptera: Acrididae)

The fine structure and distribution of various types of antennal sensilla in three nymphal stages and in adults of both solitary-reared (solitary) and crowd-reared (gregarious) phases of the desert locust, Schistocerca gregaria, were investigated by scanning and transmission electron microscopy. Four types of sensilla were identified: sensilla basiconica, s. trichodea, s. coeloconica and s. chaetica. S. basiconica contain up to 50 sensory neurons, each of which displays massive dendritic branching. The sensillar wall is penetrated by a large number of pores. In contrast, s. trichodea contain one to three sensory neurons that branch to give five or six dendrites in the sensillar lumen; the sensillum wall is penetrated by relatively few pores. The s. coeloconica are situated in spherical cuticular pits on the antennal surface. The s. coeloconica are of two types: one type contains one to three sensory neurons with double sensillar walls penetrated by slit-like pores, whereas the second type contains four sensory neurons with non-porous double sensillar walls. The s. chaetica have a flexible socket and a thick non-porous sensillum wall and contain four sensory neurons that send unbranched dendrites to a terminal pore. A fifth sensory neuron of the s. chaetica terminates in a tubular body at the base of the hair. S. basiconica and coeloconica are normally distributed over the entire antennal flagellum, with a concentration in the middle segments; s. trichodea have three areas of concentration on the 5th, 10th and 14th flagellar segments. Sensilla chaetica are most abundant on the terminal segment. Locusts raised in solitary conditions have more olfactory sensilla (s. basiconica and s. coeloconica) than crowd-reared locusts. The difference in sensillar numbers is more evident in adults than in nymphs. These results suggest that differences in the odour-mediated behaviour of nymphs and adults, and between the phases of S. gregaria, may be attributable to differences at the sensory input level.     

疟疾媒介中华按蚊触角感器的扫描电镜观察

【目的】明确中华按蚊Anopheles sinensis雌成虫与幼虫触角感器的类型、形态和分布。【方法】利用光学显微镜观察中华按蚊成虫与幼虫触角的形态结构,利用扫描电镜观察触角上的感器类型、形态和分布。【结果】中华按蚊雌成虫触角由柄节、梗节和鞭节组成,鞭节有13个亚节。触角上共发现4种类型的感器,分别为毛形感器(锐型和钝型)、刺形感器(大型和小型)、锥形感器(Ⅰ型和Ⅱ型)和腔锥形感器(大型和小型)。雌成虫触角各类感器总计约1 135.67±86.75个,其中毛形感器数量最多(662.00±6.22个),随后是刺形感器(294.67±33.35个)和锥形感器(146.00±42.39个),腔锥形感器数量最少(36.50±5.90个)。毛形感器、刺形感器和锥形感器在鞭节的每个亚节均有分布,而大型腔锥形感器在第9-11亚节没有分布,小型腔锥形感器仅分布于第13节的顶端。幼虫触角的鞭节不分亚节,呈管状,触角末端有一个感觉锥,鞭节上分布有与成虫锥形感器相似的锥形凸起,初步定名为类锥形感器,其数量和大小随幼虫龄期的增长而显著增加,锥体表面的凹槽越来越明显,其功能还需要通过超微结构和电生理等研究进一步确定。【结论】本研究对中华按蚊幼虫和雌成虫触角感器的形态特征、类型、数量及其分布进行了观察和分析,结果为进一步研究中华按蚊感器的生理功能奠定了基础。     

苹果榕传粉小蜂雌性触角感器及其生态学意义

利用扫描电子显微镜对苹果榕传粉小蜂Ceratosolen sp.雌性触角感器进行观察。结果显示:苹果榕传粉小蜂雌性触角呈膝形,分布着7类13种类型的感器,包括毛型感器、多孔的板型感器、锥型感器(2种类型)、栓锥型乳突状感器、刺型感器(4种类型)、腔锥型感器(3种类型)和一种专一性的角锥型感器。对各感器的形态、数量、分布进行描述,并结合感器的选择性染色(Ag+染色)、传粉小蜂行为及其榕/蜂互惠共生系统,对不同类型感器的功能和生态学意义进行探讨。