Comparative morphology of sensilla on antenna,maxillary palp and labial palp of larvae of Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) (Coleoptera: Curculionidae)

Eucryptorrhynchus scrobiculatus (Olivier) and E. brandti (Harold) are two wood boring pests of Ailanthus altissima (Mill.) Swingle (tree of heaven) and the variety Ailanthus altissima var. Qiantouchun. These beetles attack healthy trees and bore into roots and trunks during the larval stage. We studied the typology, distribution and morphostructure of the sensilla on the antennae, maxillary palps and labial palps of E. scrobiculatus and E. brandti larvae using scanning and transmission electron microscopy. The results showed the following: (i) the antennae of the two weevil larvae had two types of sensilla, sensilla basiconica (S.b.1 and S.b.2) and sensilla twig basiconica (S.tb.1‐S.tb.3), with S.tb.4 observed only on the antennae of E. brandti larvae; (ii) the maxillary palps had three types of sensilla, S.b.2, S.tb. (S.tb.2, S.tb.3 and S.tb.5) and digitiform sensilla; (iii) the labial palps had two types of sensilla, S.b.2 and S.tb. (S.tb.2, S.tb.3 and S.tb.5); (iv) the quantity and distribution of sensilla on the antennae, maxillary palps and labial palps remained constant between E. scrobiculatus and E. brandti larvae; and (v) sensilla basiconica had distinct sidewall pores, an apical pore was observed on sensilla twig basiconica, and digitiform sensilla were oval in shape, with a distinct apical pore. Based on the microstructure of the cuticle wall and dendrite, we hypothesized that these sensilla functioned as olfactory, gustatory and hygro‐/thermo‐receptors, respectively. We discuss the relationships among types of sensilla and the types of damage caused by the larvae inside the host tree to understand olfactory and gustatory receptor mechanisms. The results of this study will provide a firm basis for future electrophysiological studies.     

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.     

Sensilla of the antennae and the labial and maxillary palps of Blattella germanica (L.) (Dictyoptera: Blattellidae): Their classification and distribution

The various sensilla on the antennae and on the labial and maxillary palps of Blattella germanica (L.) were studied. Thick-walled chemoreceptors with fluted shafts and articulated bases are located on the antennae and on the labial and maxillary palps. Thin-walled chemoreceptors, without fluted shafts or articulated bases, are restricted to the flagellar segments of the antennae and to the distal segments of the palps. Antennae of adult males have more thin-walled chemoreceptors than do those of females. Hair-plate sensilla are found at the scape-head and scape-pedicel joints, and at the joints of segments on the palps. Campaniform sensilla are concentrated as a ring around the distal margin of the pedicel, and are also scattered singly on the scape, pedicel, and flagellar segments of the antennae, and on the first segment of the maxillary palps. Occasionally, a few sensilla coeloconica and cold receptor sensilla are found on the antennal flagellum.     

Sensing the Underground - Ultrastructure and Function of Sensory Organs in Root-Feeding Melolontha melolontha (Coleoptera: Scarabaeinae) Larvae

Introduction

Below ground orientation in insects relies mainly on olfaction and taste. The economic impact of plant root feeding scarab beetle larvae gave rise to numerous phylogenetic and ecological studies. Detailed knowledge of the sensory capacities of these larvae is nevertheless lacking. Here, we present an atlas of the sensory organs on larval head appendages of Melolontha melolontha. Our ultrastructural and electrophysiological investigations allow annotation of functions to various sensory structures.

Results

Three out of 17 ascertained sensillum types have olfactory, and 7 gustatory function. These sensillum types are unevenly distributed between antennae and palps. The most prominent chemosensory organs are antennal pore plates that in total are innervated by approximately one thousand olfactory sensory neurons grouped into functional units of three-to-four. In contrast, only two olfactory sensory neurons innervate one sensillum basiconicum on each of the palps. Gustatory sensilla chaetica dominate the apices of all head appendages, while only the palps bear thermo-/hygroreceptors. Electrophysiological responses to CO₂, an attractant for many root feeders, are exclusively observed in the antennae. Out of 54 relevant volatile compounds, various alcohols, acids, amines, esters, aldehydes, ketones and monoterpenes elicit responses in antennae and palps. All head appendages are characterized by distinct olfactory response profiles that are even enantiomer specific for some compounds.

Conclusions

Chemosensory capacities in M. melolontha larvae are as highly developed as in many adult insects. We interpret the functional sensory units underneath the antennal pore plates as cryptic sensilla placodea and suggest that these perceive a broad range of secondary plant metabolites together with CO₂. Responses to olfactory stimulation of the labial and maxillary palps indicate that typical contact chemo-sensilla have a dual gustatory and olfactory function.     

The Structure and Evolution of the Apical Sensory Zone Structures in the Maxillary and Labial Palps of Caddisflies (Trichoptera)

A comparative morphological study of the apical regions of palps in Trichoptera from different evolutionary lines of the order was carried out, and a comparison was made with representatives of Lepidoptera and Mecoptera. Light and scanning electron microscopy methods were used to study the structure of palpal apices in 81 species of the order Trichoptera, 6 of Lepidoptera, and 2 species of Mecoptera. As a result, 11 types of sensilla were identified on the terminal palpal segments, and highly differentiated apical sensory zones with thick short basiconic sensilla were found on the maxillary and labial palps. The presence of an apical sensory zone in caddisflies and related orders is recognized as a plesiomorphic character.     

Morphological and ultrastructural characterization of olfactory sensilla in Drosophila suzukii: Scanning and transmission electron microscopy

Drosophila suzukii is a serious horticultural and quarantine pest, damaging various berry crops. Although the active use of olfactory communication in D. suzukii is well-known, their olfactory sensory system has not been comprehensively reported. Therefore, the present study was carried out to understand the morphology, distribution and ultrastructure of olfactory sensilla present in the antennae and maxillary palps of D. suzukii, through scanning electron microscopy (SEM) and transmission electron microscopy (TEM). The olfactory sensilla on the antennae of D. suzukii in both sexes could be classified into three major morphological types, basiconic, trichoid and coeloconic sensilla, according to their shapes. The antennal basiconic sensilla were further divided into three subtypes and the antennal trichoid sensilla into two subtypes, respectively, according to the size of individual sensillum. In contrast to the antennal olfactory sensilla showing diverse morphology, basiconic sensilla was the only type of olfactory sensilla in the maxillary palps of D. suzukii. The basiconic sensilla in the maxillary palps could be further classified into three subtypes, based on their size. Our SEM and TEM observations indicated that multiple nanoscale pores are present on the surface of all types of olfactory sensilla in the antennae and maxillary palps, except coeloconic sensilla. The difference in the morphological types and the distribution of olfactory sensilla suggests that their olfactory functions are different between antennae and maxillary palps in D. suzukii. The results of this study provide useful information for further studies to determine the function of olfactory sensilla in D. suzukii and to understand their chemical communication system.     

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.     

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.     

Olfactory sensilla on the antenna and maxillary palp of the sheep head fly,Hydrotaea irritans (Fallen) (Diptera : Muscidae)

Antennae and maxillary palps of both sexes of the Sheep Head fly Hydrotaea irritans (Diptera : Muscidae) were investigated using scanning electron microscopy to describe the types, morphology, and distribution of olfactory sensory structures. Only socketed bristles and microtrichia were found on the scape of the antennae. These structures were also observed on the pedicel together with a group of 7–8 as yet undescribed sensilla, whose function is unknown. Olfactory sensilla were not found on these 2 segments or on the arista. Large numbers of olfactory sensilla and microtrichia were present on the funiculus. The former included sensilla trichodea (thick-walled, multiporous sensilla), sensilla styloconica and 6 types of sensilla basiconica (thin-walled, multiporous sensilla), 4 of which occurred individually and 2 of which were found in groups. An olfactory pit containing groups of thin-walled multiporous sensilla was located on the dorsomedian side of the funiculus. All sensilla basiconica were classified on morphological characteristics. The maxillary palps were covered with microtrichia and socketed bristles, but only 1 type of olfactory sensillum was found. This was a type of sensillum basiconicum that differed from any of those found on the antennae. No differences were found in sensilla diversity and distribution between males and females.     

Influence of precocene II on the sensory system of antennae and mouthparts in larvae of the fruit tree tortricid <Emphasis Type="Italic">Archips podana</Emphasis> Scop. (Lepidoptera: Tortricidae)

The influence of precocene II, an antijuvenile agent, on morphological characters of the chemoreceptor apparatus of antennae and mouthparts was studied in fifth instar A. podana larvae. Treatment with different doses of precocene was performed at the egg stage. It proved to cause changes in the form and number of basiconic sensilla on the maxillary palps and galea and in the size of basiconic sensilla on the second and third antennal segments. The results are discussed with respect to the influence of precocenes on the insect sensory system and the role of the juvenile hormone in regulation of its development.     

七星瓢虫唇须感器分布和统计特征的雌雄差异

应用扫描电镜观察七星瓢虫Coccinella septempunctata的唇须和附节,以便确认在感应农药残留时起主要作用的化学感受器。研究证明分布在下颚须上的感器类型包括两种锥形感器（兼有机械感受器和化学感受器的功能）和一种钟形感器。同时本文首次报道了对这些感器精确的形态测量,明确了其性二型性。对下颚须的测量表明其左右末节的长度差异与七星瓢虫的性别之间有明显的相关性。文中还记述了下唇须上一种新的钟形感器,其在每个下唇须上的数量大约为12～17个。经测量这些机械感受器的平均直径为2.4 μm。早期研究中所记述的锥形感器被证实是下唇须上仅有的化学感应器,并且首次证明这些锥形感器的数量与性别相关,雄性平均为18个,雌性平均为16个。附节上也存在性别差异： 第2跗节跗垫的宽度在雄性中是从后向前逐渐增大,而在雌性中刚好相反。本文还就这些差异的意义以及新发现感器的功能进行了讨论。     

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.     

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。     

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.     

Ultrastructure of sensilla on the maxillary and labial palps of three species (Coleoptera: Staphylinidae: Staphylininae)

Most species of Staphylinidae are predators in an agroecosystem. They acquire prey information from their environment through receptors. In this study, the sensilla on maxillary and labial palps of Philonthus kailiensis, Philonthus lewisius and Quedius robustus were examined with scanning electron microscopy to identify and analyse the external morphology and distribution of the sensilla to enhance our knowledge of the sensilla of Staphylinidae and provide a rationale of taxonomical studies on the two genus. Results showed that the sensilla are classified into six types: Böhm bristles, sensilla chaetica, sensilla furcate, sensilla coeloconica, sensilla placodea and sensilla basiconica. No sexual dimorphism exists among the three species. The relationships and functions of sensilla on maxillary and labial palps were also speculated. There may be a certain correlation between the sensilla on maxillary and labial palps of the staphylinid and its habitat.     

Inactivation of olfactory sensilla of a single morphological type differentially affects the response of Drosophila to odors

The olfactory organs on the head of Drosophila, antennae and maxillary palps, contain several hundred olfactory hairs, each with one or more olfactory receptor neurons. Olfactory hairs belong to one of three main morphological types, trichoid, basiconic, and coeloconic sensilla, and show characteristic spatial distribution patterns on the surface of the antenna and maxillary palps. Here we show that targeting expression of the cell-death gene reaper to basiconic sensilla (BS) causes the specific inactivation of most olfactory sensilla of this type with no detectable effect on other types of olfactory sensilla or the structure of the antennal lobe. Our data suggest that BS are required for a normal sensitivity to many odorants with a variety of chemical structures, through a wide range of concentrations. Interestingly, however, in contrast to other odorants tested, the behavioral response of ablated flies to intermediate concentrations of propionic and butyric acids is normal, suggesting the involvement of sensilla unaffected by ectopic reaper expression, probably coeloconic sensilla that respond strongly to these two organic acids. As inactivation of BS causes an underestimation of the concentration of both acids detectable at both the highest and lowest odorants concentrations, our results suggest that concentration coding for these two odorants relies on the integration of signals from different subsets of sensilla, most likely of different morphological types.     

Do grazing caddisfly larvae of Melampophylax mucoreus (Limnephilidae) use their antennae for olfactory food detection?

Abstract. The behaviour of larvae of Melampophylax mucoreus (Limnephilidae; Trichoptera) is studied to test their capacity for food detection over long distances by olfaction. In aquaria experiments with stagnant water, the larvae rapidly accumulate on a biofilm-covered stone (offered food patch) but not on control stones without biofilm-cover. Larvae in an artificial flow channel (flume), with uni-directional flow between an arrangement of stones covered with biofilm and stones without biofilm, can find food patches upstream from their location more rapidly than downstream. Stimuli transported towards the larvae by the water current might assist food location. However, no significant differences could be found in the movement of larvae with intact antennae compared with larvae without antennae. The results lead to the conclusion that the larvae of M. mucoreus find food patches mainly by random movement, which is assisted by their high mobility. Long-distance olfactory orientation towards food patches, mediated by antennae, can be precluded. Gustatory stimuli appear to play an important role in contact-chemoreception to identify biofilm patches on stone surfaces. Scanning electron microscopy studies of the larval head reveal that the antennae of M. mucoreus are small pegs (50 µm in length) with an apical plate (15 µm in diameter). The mouthparts of the larvae, especially the maxillary palps, show a variety of different sensilla types that are possibly used for contact-chemoreception. These sensilla also could be responsible for the orientation towards a food resource upstream of the larvae.     

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.     

长刺萤叶甲属与短鞘萤叶甲属的外部形态扫描电镜观察

本文对长刺萤叶甲属和短鞘萤叶甲属的头部,口器,触角,体毛,刻点,前胸背板,鞘翅缘折,足的外部形态首次进行了扫描电镜观察,并对结果进行了分析,比较,发现纬度变化引起的形态变异不显著,而海拔的变化引起形态的适应性变化比较显著。因此,对两种不同海拔的萤叶甲进行形态比较,研究萤叶甲在不同地理环境下,其形态的适应性变化提供了有价值的依据。     

Sense organs in polychaetes (Annelida)

Polychaetes possess a wide range of sensory structures. These form sense organs of several kinds, including the appendages of the head region (palps, antennae, tentacular cirri), the appendages of the trunk region and pygidium (parapodial and pygidial cirri), the nuchal organs, the dorsal organs, the lateral organs, the eyes, the photoreceptor-like sense organs, the statocysts, various kinds of pharyngeal papillae as well as structurally peculiar sensory organs of still unknown function and the apical organs of trochophore larvae. Moreover, isolated or clustered sensory cells not obviously associated with other cell types are distributed all over the body. Whereas nuchal organs are typical for polychaetes and are lacking only in a few species, all other kinds of sensory organs are restricted to certain groups of taxa or species. Some have only been described in single species till now. Sensory cells are generally bipolar sensory cells and their cell bodies are either located peripherally within the epidermis or within the central nervous system. These sensory cells are usually ciliated and different types can be disinguished. Structure, function and phylogenetic importance of the sensory structures observed in polychaetes so far are reviewed. For evaluation of the relationships of the higher taxa in Annelida palps, nuchal organs and pigmented ocelli appear to be of special importance.