Atlas of olfactory organs of Drosophila melanogaster: 1. Types,external organization,innervation and distribution of olfactory sensilla

The third antennal segment (funiculus) and the maxillary palp are the main and accessory olfactory sense organs of Drosophila melanogaster. Cryofixed antennae and palps revealed a superior preservation of the sensory dendrites and other cellular details as compared to conventional chemical fixation. Extensive cross-section series through funiculus and palp were studied in order to obtain as complete an evaluation as possible of the sensillar complement on these appendages. About 75% of all sensilla on the male and female funiculus were individually studied and their position on the antennal surface mapped. Dimensions of the cuticular apparatus of the various types of sensilla are provided as well as the number of innervating receptor neurons. Particular attention has been paid to the cuticular pores important for stimulus transport and to the sensory dendrites. On the funiculus surface, all sensilla have multiple wall pores: sensilla (s.) trichodea and s. basiconica are single-walled, s. coeloconica are double-walled. The distribution of s. trichodea and s. basiconica follows opposing gradients along a diagonal axis parallel to the axis of the arista from proximo-medial to disto-lateral. In this disto-lateral direction the density of s. trichodea increases while that of the s. basiconica decreases. S. trichodea occur in three subtypes with one, two or three receptor neurons. Basiconic sensilla can be subdivided into three subtypes of large s. basiconica (with two or four receptor neurons), three subtypes of thin s. basiconica (with mostly two, rarely four neurons), and one subtype of small s. basiconica with two receptor neurons. Large s. basiconica occur only in the most proximal region (the ‘LB-zone’); thin s. basiconica are most common in a belt that borders the LB-zone distally, while small s. basiconica are most numerous even further distally along the mentioned diagonal axis in between the s. trichodea. S. intermedia are single-walled, multiporous sensilla which combine features of s. trichodea and s. basiconica; they are found in two subtypes with two or three receptor neurons, in the same region where s. trichodea abound. The s. coeloconica are irregularly distributed over the funicular surface, and occur in two subtypes with two or three receptor neurons. Sexual dimorphism on the antenna is moderate, the female funiculus is a bit longer and carries a larger number of small s. basiconica and large s. basiconica of the LB-I subtype; the male funiculus, however, has more s. trichodea than the female. On the maxillary palp, besides mechanoreceptive s. chaetica, there are only s. basiconica with two receptor neurons. According to the fine structure of their sensory dendrites, three subtypes can be discriminated which are distributed in a random pattern. The functional significance of the described structures and distribution awaits future immunocytochemical and electrophysiological experiments.     

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.     

Structure and distribution of antennal sensilla in Oranmorpha guerinii (Gervais, 1837) (Diplopoda,Polydesmida)

Detailed information on sensory organs of Diplopoda especially on antennal sensilla are still sparse and fragmentary. The present study on the antennae of Oranmorpha guerinii (Polydesmida, Paradoxosomatidae) utilizing scanning electron microscopy revealed the presence of six sensillar types: (1) apical cones, (2) sensilla trichodea, (3) sensilla microtrichodea, (4) sensilla chaetica, (5) sensilla basiconica bacilliformia, and (6) sensilla basiconica spiniformia. External structure and distribution of cuticular antennal sensilla are compared with data from other diplopod species. We moreover discuss possible functions of antennal sensilla in millipedes.     

豆柄瘤蚜茧蜂触角感受器的扫描电镜观察(英文)

利用扫描电镜对豆柄瘤蚜茧蜂Lysiphlebus fabarum的触角感受器进行了观察, 发现有7 种感受器, 分别为毛形感受器、刺形感受器Ⅰ型、刺形感受器Ⅱ型、锥形感受器、腔锥形感受器Ⅰ型、腔锥形感受器Ⅱ型和板形感受器。毛形感受器是数量最多的感器; 除了腔锥形感受器Ⅱ型只在雌虫触角发现外, 雌、雄触角感受器的类型和结构没有较大的差别, 并且腔锥形感受器Ⅱ是数量最少的感器。     

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.     

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.     

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。     

Antennal sensilla on cave species of Australian Paratemnopteryx cockroaches (Blattaria : Blattellidae)

The external morphology and distribution of antennal sensilla of cave-dwelling Australian cockroaches, Paratemnopteryx stonei (Races B and C), P. howarthi and P. sp. nov. (Blattaria : Blattellidae), are described using scanning electron microscopy. Eight major types of sensilla were found. Long and medium-length sensilla chaetica are deeply grooved mechano and contact chemo-receptors with a terminal pore; the long type forms 5–11% and the medium-length 7–22% of all sensilla. Sensilla trichodea type 1 are very slender, non-porous, and form 43–60% of all sensilla. Sensilla trichodea type 2 are stouter, shorter and have wall pores; they form 5–14% of the sensilla. Sensilla basiconica type B are very short, non-porous, inflexible-socket receptors that are known to be hygro- and thermo-receptors; they comprise less than 0.6% of the sensilla. Sensilla basiconica type Gl are short, grooved and have a terminal pore; they form 5–11% of all sensilla. Type G2 are longer with indications of a terminal pore and form 7–10% of the sensilla. Sensilla basiconica type P are short with wall pores, and they form 3–9% of the sensilla. Total sensillar numbers ranged from 5700–8900 for P. stonei, depending on the race and sex, 6950–9950 for P. sp. nov. and 11,700-15,100 for the smaller and possibly more epigean-related P. howarthi. Females had 700–3150 fewer sensilla than males. Comparisons are made between Paratemnopteryx and common epigean cockroaches in relation to sensillar types and numbers.     

小菜蛾成虫喙管感器的超微结构及其神经元中枢投射

【目的】明确小菜蛾Plutella xylostella成虫喙管感器的形态结构及感器神经元的投射。【方法】利用扫描电子显微镜观察小菜蛾成虫喙管结构和感器,利用神经回填技术和激光共聚焦显微镜观察喙管感器神经元在脑部的投射。【结果】小菜蛾成虫喙管上存在毛形感器(两种亚型)、腔锥形感器、锥形感器、刺形感器和栓锥形感器5种不同类型的感器。毛形感器表面光滑,分布于外颚叶外侧,可分为毛形感器Ⅰ型和Ⅱ型两种亚型,其中Ⅰ型比Ⅱ型长;锥形感器分布于喙管外表面,由一个感觉锥和一个短的圆形基座组成;腔锥形感器仅分布于食管内侧,只有一个粗短感觉锥而无基座;刺形感器由一个细长的感觉毛和一个圆形基座组成,表面无孔,分布于喙管的外表面;栓锥形感器是昆虫喙管上最典型的感受器,集中分布于喙管顶端区域,感器顶部凹腔伸出一个单感觉锥。此外,喙管上的感觉和运动神经元投射到初级味觉中枢咽下神经节。【结论】本研究阐明了小菜蛾成虫喙管感器的类型、分布和形态特征及其感器神经元在脑部的投射形态,为深入了解小菜蛾喙管感器的生理和功能奠定了基础。     

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.     

Fine structure of antennal sensilla of male Aedes aegypti (L.)

The terminal two antennal segments of male Aedes aegypti bear the same variety of sensillar types as the female's antenna, namely, sensilla chaetica, sensilla coeloconica, sensilla ampullacea, grooved pegs, and four types of sensilla trichodea: long and short, pointed-tipped trichodea and blunt-tipped types I and II. Each type of sensillum has a similar fine structure in both sexes. Of the 514 neurones which innervate these sensilla in the male, 91% are olfactory receptors, 7% mechanoreceptors, and 2% thermoreceptors. The total number of neurones in the male is about four times fewer than in the female, but the ratio of those responding to the various stimuli is similar.The sensilla studied herein probably mediate stimuli involved in location of suitable resting sites and nectar meals. In addition they are apparently involved in location of vertebrates as recent studies indicate that certain male mosquitoes are attracted to hosts to bring them into the proximity of the females for mating. This host finding behaviour of males would explain why they have the same sprectrum of sensillar types as do females.     

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。     

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。     

Sensilla of the antennal flagellum in Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae)

Seven sensilla were observed on the antennal flagellum of both sexes of Pseudaletia unipuncta (Haw.) (Lepidoptera : Noctuidae): chaetica Type I and Type II), styloconica, coeloconica, trichodea (Type 1a, b, c and Type II a, b), auricillica, nonporous and squamiformia. The major sexual dimorphism noted was the presence of Type la trichodea on the male antennae, suggesting that this sensillum may contain the receptor sites for the female sex pheromone. Differences in the size of sensilla coeloconica, auricillica and trichodea (Types Ib, IIb) were noted between sexes, being larger in females. Adults obtained from larvae fed on artificial diet generally had longer antennae and sensilla than those fed on young oat plants.     

Morphological characteristics of antennal sensilla in the European cornborer Ostrinia nubilalis (Lepidoptera: Pyralidae)

The European cornborer antenna is filiform in both sexes, but exhibits a substantially larger diameter in the males. On the antenna of both sexes, the following sensillum types were characterized: sensilla trichodea, s. basiconica, s. auricillica, s. coeloconica, s. chaetica and s. styloconica. Long dorsal bristles were of a chaetic type. An intermediate trichoid/basiconic type was found in low numbers on the ventral part of the antenna. In the male, three different morphological types of s. trichodea were observed, having one, two or three sensory cells, correlated with different dimensions of the hair. The s. trichodea with three sensory cells are most common in the basal part of the antenna, while sensilla with two cells are mainly found distally. Trichodea with one sensory cell are more evenly distributed over the length of the antenna. All cells present in the different s. trichodea respond to sex pheromone components or to a behavioural antagonist in electrophysiological sensillum recordings. S. basiconica and s. auricillica had 2-3 sensory cells, and a probable olfactory function. Sensilla coeloconica, also with a putative olfactory function, contained 3-5 sensory cells. S. chaetica of the taste/tactile type possessed 4 + 1 sensory cells. S. styloconica comprised three sensory cells with possible functions as thermo- and hygroreccptors.     

Functional morphology of antennal chemoreceptors of the parasitoid Microplitis croceipes (Hymenoptera: Braconidae)

The specialist parasitoid, Microplitis croceipes (Cresson) (Hymenoptera: Braconidae), uses chemical cues from plants damaged by herbivore-feeding and also plant by-products in host location and acceptance. These chemicals are detected by the wasp's antennae. We conducted scanning and transmission electron microscopy studies of male and female M. croceipes antennae and detected no distinct morphological differences in the chemoreceptors between the sexes. Male antennae are approximately twice as long as female antennae. We found five morphological sensillar types in both sexes: sensilla (s) trichodea were the most abundant and distributed over the whole antenna; s. chaetica were present in low numbers only on the scape and pedicel; and s. basiconica, s. coeloconica and elongated s. placodea were found only on the flagellum. Ultrastructural investigations revealed pore systems on s. basiconica and s. placodea. In s. placodea, sensory neurons run parallel to the longitudinal axis of the sensilla with terminal pores. We recorded responses from single olfactory receptor neurons in s. placodea to two plant-emitted volatiles, cis-3-hexenol and ocimene, and two anthropogenic compounds, cyclohexanone and 2-diisopropylaminoethanol. Male receptor neurons were more sensitive than those of females with significantly higher spike frequency being registered from neurons in males.     

The antennae and antennal sensilla of the giant looper,Boarmia (Ascotis) selenaria SchiffermÜller (Lepidoptera,Geometridae)*

The general morphology of the antennae and the antennal sensilla of both sexes of Boarmia (Ascotis) selenaria SchiffermÜller are described. The following types of sensilla were found: Bohm bristles and sensilla chaetica, trichodea, coeloconica, auricillica and styloconica. Antennal sexual dimorphism is marked, but mainly confined to the presence of long sensilla trichodea in males and the absence in females. The sensilla trichodea are the most abundant sensillum type.     

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.