山茱萸蛀果蛾幼虫触角和口器感器的超微形态

【目的】探索寄主范围不同的蛀果蛾科(Carposinidae)幼虫感器之间是否存在差异。【方法】采用扫描电子显微镜观察了为害山茱萸Cornus officinalis Sieb. et Zucc.的单食性蛀果害虫--山茱萸蛀果蛾Carposina coreana Kim老熟幼虫触角和口器感器的超微形态。【结果】山茱萸蛀果蛾幼虫触角柄节未见感器分布, 梗节上有2个刺形感器和3个锥形感器, 鞭节上有1个栓锥感器和3个锥形感器。口器上共有6种感器： 刺形感器数量多, 分布广; 栓锥感器主要分布在颚叶、 下颚须和下唇须上; 指形感器位于内唇和下颚须端节侧缘; 锥形感器和板形感器仅存在于下颚上; 内唇感器为内唇所特有。【结论】蛀果蛾幼虫触角和口器的感器与寄主范围之间未发现严格的对应关系。     

Morphology,distribution, ultrastructure,and possible function of the external sensilla on antennae and mouthparts of Echinothrips americanus Morgan (Thysanoptera: Thripidae)

To better understand the semiochemical detection and reception mechanisms that involved in intra- and interspecific communication, host selection and mating behavior of Echinothrips americanus (Thysanoptera: Thripidae), scanning electron microscopy was used to examine the external morphology, distribution and ultrastructure of the sensilla on antennae and mouthparts. According to serial SEM images, seven types of antennal sensilla in total were observed in both sexes. Sensilla on mouthparts were observed concentrating on the maxillary palps, labial palps, and paraglossa, which is consistent with the previous conclusions. Possible function of the external sensilla is also discussed.     

中华蚊蝎蛉成虫口器感器的超微结构

口器感器在昆虫取食活动中起着重要作用, 但蚊蝎蛉成虫口器上感器的种类和形态迄今未见报道。我们利用扫描电子显微镜, 观察了中华蚊蝎蛉Bittacus sinensis Walker成虫口器上的感器。结果显示: 中华蚊蝎蛉口器上共有8种感器, 分别为锥形、毛形、刺形、指形、掌状、钟形、柱状感器及Böhm氏鬃毛, 主要集中于内唇、 下颚须以及下唇须上。锥形感器和刺形感器数量最多; 毛形感器主要在下颚轴节、 茎节和下唇的亚颏和前颏有分布; 钟形感器和Böhm氏鬃毛只存在于下唇须和下颚须上。下颚须端节和下唇须端节的感器种类相同, 以锥形感器为主。高度骨化的上颚以及下颚内颚叶与外颚叶上未发现感器分布。简要讨论了口器感器在昆虫分类中的意义。     

Ultrastructure of sensilla of the female mountain pine beetle,Dendroctonus ponderosae hopkins (Coleoptera : scolytidae)

The objective of this study is to provide a morphological foundation for electrophysiological studies on the sensilla of D. ponderosae. The sensilla on the antennae, labial and maxillary palps, galeae and fore-tarsi are described. The antennal club has several types of sensilla: (1) multiporous non-socketed pegs of varying lengths innervated by 2 neurons, (2) uniporous socketed pegs innervated by 5 neurons, (3) pegs innervated by only 1 neuron at the base, (4) tapered uniporous cones innervated by 4 neurons, and (5) fluted multiporous, non-socketed cones innervated by 4 neurons. All hairs on the fore-tarsi are innervated by 1 neuron at the base. The labial and maxillary palp-tips have: (1) digitiform organs, (2) campaniform organs, (3) uniporous, socketed pegs innervated by 5 neurons, and (4) non-porous pegs innervated by 3 neurons. Two small, uniporous, socketed pegs innervated by 4 neurons, are set into the maxillary palp-tip sidewall, and a single, sharp-tipped peg, innervated by 1 neuron, is found on the tip. These 2 types are not seen on the labial palps. The ultrastructure of each type of sensillum found on both the maxillary and labial palps is similar; however, the number of uniporous and non-porous pegs on the labial palps is about half those of the maxilla. The maxillary galeae each have 3 types of sensilla: (1) numerous pegs of various sizes innervated by 1 neuron, (2) a single, possibly uniporous socketed peg innervated by 5 neurons, and (3) another single non-porous non-socketed peg innervated by 2 neurons.     

小菜蛾成虫下唇须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成虫下唇须感器的形态结构及感器神经元的投射。【方法】利用光学显微镜观察和扫描电子显微镜观察下唇须结构和感器类型,利用神经回填技术和激光共聚焦显微镜观察下唇须感器神经元在脑部的投射。【结果】小菜蛾成虫下唇须共3节,其上存在Böhm氏鬃毛、钟形感器、鳞形感器、锥形感器、微毛形感器5种不同类型的感器和一个陷窝器结构。Böhm氏鬃毛短小尖细,钟形感器形如顶部凹陷的圆帽,两种感器均分布于下唇须第1节,且大小上均无雌雄二型差异;鳞形感器形同柳叶,锥形感器粗而直,均散生于下唇须的第2和3节,两种感器在大小上均存在雌雄二型差异,其中雌性的鳞形感器显著大于雄性的,根据其雌雄二型差异现象推测雌蛾的鳞形感器可能与感受寄主植物挥发物有关;下唇须第3节中上部具有一个圆形陷窝器结构,雄虫的陷窝器内径为5.68±0.33μm,雌虫的为6.03±0.23μm,雌雄间无显著性差异;凹坑内长有表面光滑的微毛形感器。小菜蛾下唇须感器神经元主要投射于脑部咽下神经节、每个触角叶的下唇须陷窝器神经纤维球和腹神经索3条通路。【结论】阐明了小菜蛾下唇须感器的类型、分布和形态特征及其感器神经元在脑部的投射形态,为深入了解小菜蛾下唇须感器的生理和功能奠定了基础。     

Maxillary appendages used by western corn rootworms, Diabrotica virgifera virgifera, to discriminate between a phagostimulant and -deterrent

Ablation of maxillary appendages on western corn rootworm, Diabrotica virgifera virgifera LeConte (Coleoptera: Chrysomelidae), were used to determine which mouthparts were involved in discrimination between the phagostimulant, cucurbitacin B, and the phagodeterrent, strychnine. Strychnine deterred feeding in unablated beetles when its concentration on disks was equal to or higher than cucurbitacin B. Unilateral or bilateral lacinea ablation did not affect discrimination between cucurbitacin B and strychnine. Galeal ablations abolished strychnine deterrence at equimolar concentrations of both compounds but not at higher strychnine concentrations, whereas maxillary palp ablation had no effect. However, bilateral ablation of both galea and maxillary palps resulted in a loss of deterrence at all strychnine concentrations. These results indicated strychnine sensitive chemoreceptors were primarily located on the galeae and maxillary palps. However, beetles without galeae and palps still feed on disks suggesting that cucurbitacin B sensitive chemoreceptors were located on other mouthparts and contributed to cucurbitacin B phagostimulation.     

Taste receptors in Colorado beetle larvae

Electrophysiological studies show that at least six sensilla on the various mouthparts of larval Colorado potato beetles (Leptinotarsa decemlineata) contain chemosensory cells. These sensilla are found on the galea, maxillary palps, and labial palps. Sodium chloride, sucrose, some amino acids, and chlorogenic acid are among the effective stimuli. The sensitivity spectrum of these cells correlates with known larval phagostimulants.     

锈翅蚁蛉头部附器感器的扫描电镜观察

利用扫描电镜对锈翅蚁蛉Myrmeleon ferrugineipennis Bao&Wang雌雄成虫头部触角及口器感器的形态进行观察,描述了感器的种类、数量和分布,以期解析其取食机制。结果表明:锈翅蚁蛉触角上存在10种感器,即毛形感器、锥形感器、刺形感器、腔形感器、钟状感器、鳃形感器、耳形感器、盘形感器、舌形感器和Bhm氏鬃毛,其中毛形感器有3种亚型,数量最多;耳形感器、腔形感器和钟状感器仅在雌成虫触角上发现,而舌形感器和鳃形感器仅在雄成虫触角上发现;在锈翅蚁蛉触角鞭节近末端扁平匙状处各有1枚盘形感器,其形状和位置在雌雄虫上有差异。鳃形感器和盘形感器在已有的昆虫感器研究中未见报道,是新发现的昆虫触角感器。下颚须、下唇须上均发现锥形感器,下唇须上的数量多于下颚须;此外,下颚须上还存在钟状感器。     

Sensory Organs in Ips typographus (Insecta: Coleoptera) Fine structure of the sensilla of the maxillary and labial palps*

The labial and maxillary palps of the bark beetle, Ips typographus, possess a diversified array of sensilla. There are four types of sensilla possessing a single tubular body indicating a mechanoreceptive function. The variation of the associated cuticular structures of these sensillar types ranges from long bristles to cavities within the cuticle. Terminal pore sensilla with a supposed mechanosensory/gustatory function and single-walled presumably chemoreceptive sensilla are found on the apical part of the terminal palp segments. A poreless sensillar type is found on the lateral side of the terminal segment of the maxillary palp. The functional capabilities of this sensillar type are presently unknown.     

Sensilla on the larval antennae and mouthparts of the european sunflower moth,Homoeosoma nebulella Den. and Schiff. (Lepidoptera : Pyralidae)

Sensilla on the larval antennae and mouthparts of Homoeosoma nebulella (Lepidoptera : Pyralidae) were examined with a scanning electron microscope. The antenna possesses 2 aporous sensilla chaetica, one aporous sensillum styloconicum, 3 large multiporous sensilla basiconica, and 3 small multiporous s. basiconica. The labrum bears 12 aporous s. chaetica and the epipharynx has 6 aporous flattened s. chaetica and 4 epipharyngeal sensilla. It is the first time that more than 2 epipharyngeal sensilla have been reported in caterpillars. The mandible possesses 4 aporous s. chaetica. Each galea bears 2 uniporous s. styloconica, 2 small aporous s. basiconica, one s. campaniformium and 3 aporous s. chaetica. Each maxillary palp bears a distal group of 8 uniporous s. basiconica, one large s. digitiformium and one pore plate. Minute grooves exist on the s. digitiformium. Each labial palp possesses one aporous s. chaeticum and one aporous s. styloconicum. An attempt has been made to identify the function of sensilla by comparing them with those of other larvae.     

Sexual dimorphism of adult labial palps of the peach fruit moth Carposina sasakii Matsumura (Lepidoptera: Carposinidae) with notes on their sensilla

The labial palps and their sensilla of the peach fruit moth Carposina sasakii Matsumura, a serious pest of pome fruits in eastern Asia, were investigated using light microscopy and scanning and transmission electron microscopy. The labial palps are three‐segmented and exhibit distinct sexual dimorphism in length: much longer in the female than in the male. Four types of sensilla were found on the labial palps: campaniform, squamiform, chaetic and flattened sensilla. The campaniform sensilla are present on the first segment alone. The squamiform sensilla are located on all the three segments. The chaetic sensilla are mainly present on the third segment. The flattened sensilla are grooved with wall pores and situated in a labial palp‐pit organ, which is located at the apex of the distal segment of the labial palp. The sexual dimorphism of the labial palp and the labial palp‐pit organ was briefly discussed.     

Morphology of antennal,maxillary palp and labial palp sensilla in different larval instars of the Asian long‐horned beetle,Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae)

The Asian long‐horned beetle (ALB) is one of the most important international trunk‐boring insect pests of forest trees, which has caused enormous losses in the forests of China, the USA and some countries in Europe. To investigate the morphology of the antennal, maxillary palp and labial palp sensilla in the five larval instars of the ALB, scanning electron microscopy was used to determine the typology, morphology, number and distribution of the sensilla. The antennae had two types of sensilla: the sensilla twig basiconica (s.t.b.) and sensilla basiconica. Three different types of sensilla were observed on the maxillary palps, which were the sensilla digitiformia, the sensilla chaetica and the s.t.b. The labial palps had two types of sensilla: the sensilla chaetica and the s.t.b. The s.t.b. had seven total subtypes, and the sensilla basiconica had two subtypes. The typology, quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant in the five larval instars, but the dimensions of all sensilla types increased throughout the development. We discussed the mechanoreception and the chemoreception of ALB larvae inside the host tree to provide a theoretical understanding and information for further research on the behaviour and the electrophysiology of this devastating forest pest.     

Ultrastructure and distribution of sensilla on the maxillary and labial palps of Chlorophorus caragana (Coleoptera: Cerambycidae)

Chlorophorus caragana is a species of long‐horned beetle that damages Caragana davazamcii Sancz. (Fabales: Papilionaceae) bushes in desert areas in China. The beetles cause substantial damage to local forestry plantations and the environment. Sensilla on the maxillary and labial palps of coleopterans a allow the insects to recognize their host plants. We used scanning and transmission electron microscopy to study the ultrastructure, distribution, and abundance of various sensilla on the maxillary and labial palps of C. caragana. We found four types of sensilla including ten subtypes: one of Böhm's bristles, three of sensilla chaetica, one of digitiform sensilla, and five of sensilla twig basiconica. The types and distribution of the sensilla on the maxillary and labial palps were highly similar between males and females. Finally, this article discusses the functions of the sensilla of related species in recognizing hosts and the significance of gustation studies in the context of the control of C. caragana.     

Sensory organs of the antennae and mouthparts of beetle larvae (Coleoptera)

The sensilla located on the antennae and maxillary and labial palps of the larvae of 64 beetle species from 22 families were studied using electron microscopy. The larvae of beetles living in different habitats and having different trophic specializations possess a uniform structure of the sensory organs. They are composed of two groups of sensilla on the apical and subapical segments of the antennae, one apical group of sensilla on both maxillary and labial palps, and one or several digitiform sensilla on the lateral surface of the maxillary and, occasionally, labial palp. The external morphology of the sensory organs is adaptive and represents modifications of the initial type. Band-shaped sensilla or placoid sensilla, clearly different from the initial sensory organs, appear in some taxa as rare exceptions, while other groups display either partial reduction of the receptor organs (Gyrinidae) or reduction of the cuticular parts of the sensilla (Cantharidae).     

Fine structure and functional morphology of the mouthparts of Bittacus planus and Terrobittacus implicatus (Insecta: Mecoptera: Bittacidae)

The Bittacidae are unique in Mecoptera for their adults being predaceous. However, their mouthparts have not been well documented for functional morphology to date. Here, we investigated the mouthpart morphology of the hangingflies Bittacus planus Cheng and Terrobittacus implicatus (Huang & Hua) using scanning electron microscopy. The mouthparts are of the mandibulate type and situated at the tip of an elongated rostrum. The labrum is greatly elongated, roughly twice as long as the subquadrate clypeus. The epipharynx is furnished with a row of basiconic sensilla arranged evenly as a median band extending from the apex to the base. The mandibles are slender and elongated, bearing a sharp lateral and a small mesal tooth. The maxillae are well developed, each consisting of a partially sclerotized cardo and a stipes, a hirsute galea and a lacinia, and a five-segmented maxillary palp. The sensillar pattern on the distal segment of the maxillary palp differs slightly between the two bittacid species. The labium is composed of a postmentum, a prementum, and a pair of two-segmented labial palps. The feeding mechanism of bittacids is briefly discussed in combination with the mouthpart morphology and their feeding habits.     

Functional Morphology and Sexual Dimorphism of Mouthparts of the Short-Faced Scorpionfly Panorpodes kuandianensis (Mecoptera: Panorpodidae)

Mouthparts are closely associated with the feeding behavior and feeding habits of insects. The features of mouthparts frequently provide important traits for evolutionary biologists and systematists. The short-faced scorpionflies (Panorpodidae) are distinctly different from other families of Mecoptera by their extremely short rostrum. However, their feeding habits are largely unknown so far. In this study, the mouthpart morphology of Panorpodes kuandianensis Zhong et al., 2011 was investigated using scanning electron microscopy and histological techniques. The mandibulate mouthparts are situated at the tip of the short rostrum. The clypeus and labrum are short and lack distinct demarcation between them. The epipharynx is furnished with sublateral and median sensilla patches. The blade-shaped mandibles are sclerotized and symmetrical, bearing apical teeth and serrate inner margins. The maxilla and labium retain the structures of the typical pattern of biting insects. The hirsute galea, triangular pyramid-shaped lacinia, and labial palps are described in detail at ultrastructural level for the first time. Abundant sensilla are distributed on the surface of maxillary and labial palps. The sexual dimorphism of mouthparts is found in Panorpodes for the first time, mainly exhibiting on the emargination of the labrum and apical teeth of mandibles. Based on the features of mouthparts, the potential feeding strategy and feeding mechanism are briefly discussed in Panorpodes.     

Fine structure and primary sensory projections of sensilla on the maxillary palp of Drosophila melanogaster Meigen (Diptera : Drosophilidae)

Three types of hairs were identified on the maxillary palp of Drosophila melanogaster Meigen (Diptera : Drosophilidae): (i) single-walled, multiporous sensilla basiconica, which constitute 75% of the innervated hairs; (ii) thick walled non-porous sensilla trichodea, which make up the remaining 25% of the innervated hairs; and (iii) numerous spinules, which are un-innervated. These sensilla basiconica uniformly contain 2 bipolar sense cells, whereas sensilla trichodea have a single dendrite with a tubular body at the base of each hair. A majority of the sensilla basiconica is located on the distal half of the dorsal surface, whereas sensilla trichodea are positioned on the tip and entire ventrolateral ridge of the palp. Approximately 125 axons of the sense cells join to form a single nerve. The structure of sensilla basiconica and sensilla trichodea suggests that they are olfactory and mechanosensory respectively. The contact chemoreceptors (gustatory sensilla) are conspicuously absent on the maxillary palp.Golgi silver impregnations and cobalt fills show that the primary sensory fibres from sensilla trichodea and sensilla basiconica on the maxillary palp project in the posterior suboesophageal ganglion (SOG) and the antennal lobe respectively. A single fibre projects separately either in the SOG or in the antennal lobe. In the antennal lobe, the input received from sensilla basiconica is usually bilateral and at least 5 glomeruli are innervated symmetrically on either side from both the palps.This study suggests that the sensory neurons are capable of making selective projections in the specific regions of the brain. Accordingly, the fibres from a sensillum project to the brain with respect to their functions and the individual glomeruli represent functional units of the brain, receiving inputs in a characteristic combination.     

Effects of juvenile hormone on sense organs involved in mating behaviour of Blattella germanica (L.) (Dictyoptera: Blattellidae)

Possible causes of the suppression of mating in juvenile hormone-treated Blattella germanica females, including the roles of the compound eyes and antennal, labial and maxillary palp sensilla in mating, were studied.Either antennectomy or labial palpectomy, but not maxillary palpectomy, leads to complete suppression of mating in females. Vision plays no role in mating. These results show that all the head appendages (antennae and labial and maxillary palps) are important in precopulatory behaviour, individually and collectively. Juvenile hormone treatment of the sixth (last) instar female nymphs leads to retention of nymphal sensillar characteristics on both the antennae and the palps of the resulting adults, with a reduced electroantennogram response. Thus the animal behaves like a nymph and does not mate in spite of its ability to produce pheromone.     

Sensilla on maxillary and labial palps in a helicophagous ground beetle larva (Coleoptera,Carabidae)

Sensilla on the labial and maxillary palp of Carabus lefebvrei Dejean, 1826 larvae were investigated using scanning and transmission electron microscopy. Two identical sets of sensilla were present on the tips of both palp pairs, and four morphological types of sensilla were identified: sensilla basiconica types 1 and 2, sensilla coeloconica and sensilla digitiformia. Ultrastructure indicates that the sensilla basiconica type 1 and coeloconica have a chemical role as gustatory and olfactory receptors, respectively, while sensilla basiconica type 2 are mechanoreceptors and the sensilla digitiformia are probably thermo‐ and hygroreceptors. Their function is discussed in relation to specialized prey detection and habitat adaptations.     

Ultrastructure and Comparative Morphology of Mouth-part Sensilla in Ground Beetle Larvae (Insecta, Coleoptera, Carabidae)

Labial and maxillary palps in 22 species of ground beetle larvae revealed the presence of four different types of sensilla: sensilla digitiformia, sensilla campaniformia, sensilla basiconica and sensilla chaetica. Using transmission electron microscopy techniques, the role of most of them was defined: sensilla campaniformia and sensilla chaetica are mechanoreceptors; while sensilla basiconica are chemoreceptors. Sensilla digitiformia can be both mechanoreceptors and chemoreceptors, though hygroreception may also be postulated on the basis of their abundance on the mouthparts of some hygrophilous species that depend on damp or wet patchy biotopes/seasons. The number and location of the sensory structures has been found to vary among the investigated taxa. The differences are related to the larval behavioural types, and are an adaptive response to different lifestyles, feeding habits and use of space.