Fine structure of olfactory sensilla in myriapods and arachnids.

Structural features of various types of olfactory sensilla are reviewed. 1) Sensilla basiconica which differ in form and size are found on the antennae of centipedes and millipedes. Their walls show longitudinal slits or grooves that either open into the sensillum lumen or do not penetrate the cuticle. In other such sensilla the outer surface is pierced by pores and the inner surface grooved and pocketed. These sensilla are innervated by one to six sensory cells. Their unbranched outer dendritic segments extend to the tip of the sensillum. The sensory cells are surrounded by two or three sheath cells which terminate at the sensillum base or form a continuous tube around the entire length of the outer dendritic segments. 2) Temporal organs of centipedes are located between the insertion of the antenna and the ocelli. These sensilla consist of a shallow cuticular ring with a central sensory plate made up by a layer of unperforated cuticle or a capsule with a mushroom-shaped structure inside formed by fibrous-looking cuticle. A dozen sensory cells with unbranched outer dendritic segments innervate each sensillum. They extend toward the sensory cuticle and pass just below it. Numerous sheath cell processes run parallel to the outer dendritic segments up to the sensory cuticle. 3) Thread-like flagella of Pauropoda are found on the antennae. They possess a flexible unperforated cuticular wall. These sensilla contain nine sensory cells surrounded by several sheath cells which form a continuous cytoplasmic tube around the outer dendritic segments. 4) Single-walled sensilla with numerous plugged pores penetrating the cuticular wall occur on the tarsus of the first leg in ticks. Each sensillum is innervated by 4-15 sensory cells. Three sheath cells terminate in the base of the sensillum. 5) Double-walled sensilla with spoke canals are found on the first tarsus of ticks. Their shaft is longitudinally grooved. Pore canals lead inward from the bottom of the grooves and open into vase-shaped chambers. From its base these canals extend into the lumen of the sensillum which contains unbranched outer dendritic segments of 1-2 sensory cells. 6) Single-walled sensilla with pore openings occur on the distal tarsal segments of the first leg of whip spiders. These sensilla are innervated by 40-45 sensory cells. Their unbranched outer dendritic segments fill the shaft lumen and extend partly into the wall pores. Microvillus-shaped sheath cell processes line the inner surface of the cuticular wall.(ABSTRACT TRUNCATED AT 400 WORDS)     

Fine structure of a developing insect olfactory organ: morphogenesis of the silkmoth antenna.

The olfactory organ of the silkmoth Antheraea polyphemus is the feathered antenna which carries about 70,000 olfactory sensilla in the male. It develops within 3 weeks from a leaf-shaped epidermal sac by means of segmental primary and secondary indentations which proceed from the periphery towards the centerline. During the first day post-apolysis, the antennal epidermis differentiates into segmentally arranged, alternating sensillogenic and non-sensillogenic regions. Within the first 2 days post-apolysis, the anlagen of olfactory sensilla arise from electron-dense mother cells in the sensillogenic epidermis. The axons of the developing sensilla begin to form the primary innervation pattern during the second day. The sensilla develop approximately within the first 10 days to their final shape, while the indentations are completed during the same period of time. The indentations are most probably driven by long basal extensions of epidermal cells, the epidermal feet. Primary indentations follow the course of segmentally arranged tracheal bundles and form the segments of the antenna. The secondary indentations follow the course of the primary segmental nerves which are reconstructed by this process. During the remaining time of development, the cuticle of the antenna and the sensory hairs is secreted by the epidermal and the hair-forming cells.     

The aesthetasc concept: structural variations of putative olfactory receptor cell complexes in Crustacea.

The structure of the aesthetascs has been investigated in the prawn Macrobrachium rosenbergii (larvae and juveniles), the opossum shrimp Neomysis integer, the euphausid Meganyctiphanes, and in the water-fleas Daphnia magna and D. longispina. The aesthetascs, that are thought to represent olfactory receptors, exhibit a considerable structural variation, ranging from the well known aesthetascs of higher crustaceans (lobster, crab, crayfish) to the corresponding sensilla found in the water-fleas and the males of opossum shrimps. The two following morphological characteristics of the aesthetascs are thought to indicate an olfactory function: the shape of the cuticular hair that is long and essentially hose-shaped, and the thin, loosely arranged cuticle of at least the outer part of the cuticular hair. The presence of other structural elements such as sensory cells, cilia, and enveloping cells are vital for the olfactory function, but the development is variable, which makes their use in the morphological definition of aesthetascs problematic.     

Antennal and mouthpart sensilla of Acanthoscelides obtectus say (Coleoptera: Bruchidae)

Acanthoscelides obtectus, a serious pest with a worldwide distribution, damages stored seeds of Phaseolus sp. To acquire a better understanding of the sensory orientation of bean weevils, their antennal and palpal sensilla were investigated. The antennae and palps possess several types of sensory hairs: blunt‐tipped sensilla chaetica and trichoidea and also sharp‐tipped sensilla trichoidea, these last being the most numerous. Sensilla basiconica are present only on the flagellomeres, but Böhm bristles occur on the basal sclerite and scape of the antennae. The terminal segments of the maxillary and labial palps bear a group of sensilla styloconica with single apical pores. Sensilla coeloconica are located only on the terminal palpal segments. They were unable to detect any distinct morphological differences between males and females. The possible chemosensory, mechanosensory, and thermo‐/hygroreceptive functions of the sensilla are discussed.     

Physiological significance of gravity receptors on larval cephalic cuticle in the silk moth,Antheraea proylei Jolly

The lateral aspects of larval cephalic cuticle of oak tasar moth, Antheraea proylei, a hybrid between Antheraea pernyi and Antheraea roylei exhibited the presence of gravity receptors in the form of dorsal campaniform sensilla. The distribution pattern and number of dome shaped dorsal campaniform sensilla were found to vary in different larval stages. In the first and second larval stages, 4‐5 sensilla were localized near the apex of the lateral aspects of the cephalic cuticle on either side of the head. From the third larval stage onwards, on the other hand, both the right and left lateral aspects of the cephalic cuticle were covered with innumerable dome shaped dorsal campaniform sensilla. The sensilla were found to be arranged in groups of 3 to 5 on the cephalic cuticle with dome‐free cuticular portion of about 50–100 µm in length between two adjacent groups of the sensilla. The individual dome shaped dorsal campaniform sensillum was either smooth surfaced or was covered with smaller domes through out the surface. The significance of differences in the number and distribution pattern of the dorsal campaniform sensilla among different larval stages in relation to gravity reception and preferred feeding posture are discussed in the light of available literature.     

Post-embryonic development of the antennal sensilla in Melipona quadrifasciata anthidioides (Hymenoptera: Meliponini)

The sensilla are sensory organs formed by cuticular and cellular structures specialized in reception of chemical and physical stimuli from the environment and transmission to the insect's central nervous system. In function of the great concentration of sensilla, the antennae are the main organs for interaction between bees and with the environment. This work studied the presence of antennal sensilla in the different phases of pupal development of the stingless bee Melipona quadrifasciata anthidioides by means of scanning electron microscopy and light microscopy. The results showed that antennal sensilla begin their development in the transition of the prepupae to the white-eyed pupae and finish it in the pigmented-body pupae phase. The antennal sensilla were exposed to the environment in the black-eyed pupae when the old cuticle is completely digested, suggesting that only in the final pupal phases can these bees perceive the environmental stimuli.     

Larval antennal sensilla in water-living insects

An overview of larval antennal sensilla in hemimetabolous and holometabolous water-living insects is given by updating current knowledge on the fine structure of these sensory systems. In the absence of successful electrophysiological studies, the possible function of sensilla is deduced from their architecture. Various kinds of sensilla are described in hemimetabolous insects, such as Ephemeroptera, Odonata, and Plecoptera, and holometabolous insects, such as Diptera Nematocera (Culicidae, Simuliidae, Psychodidae, Chaoboridae) and Trichoptera. Their possible function in responding to stimuli from the freshwater environment is illustrated and discussed. The importance of sensilla as taxonomic and phylogenetic traits is reported in Baetidae (Ephemeroptera) and in Diptera Nematocera. Some homologies outlined in Diptera Nematocera give evidence of a link between species, such as Chaoboridae and Psychodidae, whose antennae are highly modified, prehensile in the former and very reduced in the latter. Particular features, such as cuticle without pores in chemosensory sensilla and naked perikarya, are so far found exclusively in some water-living arthropods, thus reflecting a possible adaptation to the aquatic habitat. The structure of sensilla and chloride cells, which have a similar external morphology, is presented and discussed in various insect groups, considering the possible derivation of the chloride cells from sensilla.     

Surface ultrastructure of larval mouthpart sensilla of the muga silkmoth,Antheraea assamensis,an endemic species of North‐East India

Scanning electron microscopy of different larval stages of the muga silk moth Antheraea assamensis revealed the presence of sensilla chaetica, sensilla trichoidea, sensilla styloconica, gustatory sensilla, sensory pegs, placoid sensilla, etc., on their mouth parts. Some variations were observed in the number and sub‐types of sensilla in certain larval stages indicating some differences in sensitivity of the worms in different instars to the food plant and microhabitat. Microsc. Res. Tech., 2010. © 2010 Wiley‐Liss, Inc.     

Distribution patterns and morphology of sensilla on the apical segment of the antennae and palpi of Hydradephaga (Coleoptera: adephaga).

The sensilla on the terminal antennomere of selected Hydradephagans (Coleoptera) are modified in several distinct ways as compared to the sensilla on the terminal antennomere of Geadephagans (Coleoptera). There are no long sensilla of any type on antennae of Hydradephagans, the sensilla are either short, peg-like sensilla that may be recessed in the antennal surface or they are multiporous plate-like sensilla. Multiporous plate-like sensilla have not been found on the antennae of Geadephagans. A pit sensillum occurs on the antennae of one family, Gyrinidae, in the Hydradephaga whereas it is found on the antennae of several species from the families of Geadephaga. The sensilla on the terminal antennomere of Hydradephaga are often grouped into sensory fields that could be located on a particular area of the antennomere. Sensory fields are also located on the apices of the maxillary and labial palps and the most unique sensory fields occur in the Noteridae. The maxillary palpal apex of Hydrocanthus oblongus (Noteridae) is bifurcate and a sensory field with several types of sensilla is at the apex of each branch. The terminal labial palpomere is greatly enlarged and has a long, slender sensory field at the apex and a secondary sensory field on a thumb-like projection. The sensilla on the palpal apices are more diverse and complex in form as compared to the sensilla on the palps of Geadephaga. The most complex sensillum has a cuticular collar and a dome that is divided into six or seven triangular sections. The dome may be opened or closed and a peg-like sensillum with a very wide apical pore is situated beneath the dome.     

Sensilla and secretory glands in the antennae of a primitive ant: Dinoponera lucida (Formicidae: Ponerinae)

Morphology of the antennae of the female workers of the ponerine ant Dinoponera lucida was examined by light and scanning electron microscopy. In several antennomers, we found secretory gland cells of class I and III. Class III gland cells release their secretion through single pores in the antennal surface, whereas class I secretory cells are seen as tall epidermal cells close to the cuticle. Both gland types have weak reaction for total proteins and neutral polysaccharides. Six distinct sensilla types were observed: trichodea, chaetica, campaniform, basiconica, placodea, and coeloconica. The possible sensory functions of these sensilla and the gland functions are discussed.     

Morphology,distribution, and abundance of antennal sensilla of male and female macropterous and brachypterous small brown planthopper,Laodelphax striatellus (Fallén) (Hemiptera: Delphacidae)

The antennal sensilla of both genders of macropterous and brachypterous adults of the small brown planthopper, Laodelphax striatellus (Fallén) (Hemiptera: Delphacidae) were examined using light and scanning electron microscopy. Scanning electron microscopy revealed seven types of antennal sensilla in adult L. striatellus which were not evenly distributed on all antennal segments. Sensilla chaetica, a sensillum campaniformium and a Böhm bristle were found on the scape. Sensilla chaetica, sensilla trichodea, sensilla placodea which always present as plaque organs, sensilla basiconica and a sensillum campaniformium were present on the pedicel. Three sensilla basiconica and one sensillum coeloconicum containing two sensory pegs were located on the swollen sensory region of the basal flagellum. Pores observed on the surface of s. trichodea and s. placodea suggest these organs probably play a role in olfaction, whereas the aporous s. chaetica with flexible sockets probably function as mechanoreceptors. The aporous s. basiconica with inflexible sockets are probable to be thermo‐hygroreceptors while the Böhm bristle and s. campaniformia may act as antennal proprioceptors. The function of s. coeloconicum remains uncertain. The numerical dominance of antennal olfactory receptors suggests olfaction is an important function of the antenna in L. striatellus. Although a small degree of sexual/wing dimorphism was observed in the numbers of sensilla and in the length and width of antennae and antennal segments, the basic shape and structure of the antennae and antennal sensilla did not differ between the gender or wing form in L. striatellus. Microsc. Res. Tech. 2012. © 2012 Wiley Periodicals, Inc.     

Peripheral synaptic contacts at mechanoreceptors in arachnids and crustaceans: morphological and immunocytochemical characteristics

Two types of sensory organs in crustaceans and arachnids, the various mechanoreceptors of spiders and the crustacean muscle receptor organs (MRO), receive extensive efferent synaptic innervation in the periphery. Although the two sensory systems are quite different-the MRO is a muscle stretch receptor while most spider mechanoreceptors are cuticular sensilla-this innervation exhibits marked similarities. Detailed ultrastructural investigations of the synaptic contacts along the mechanosensitive neurons of a spider slit sense organ reveal four important features, all having remarkable resemblances to the synaptic innervation at the MRO: (1) The mechanosensory neurons are accompanied by several fine fibers of central origin, which are presynaptic upon the mechanoreceptors. Efferent control of sensory function has only recently been confirmed electrophysiologically for the peripheral innervation of spider slit sensilla. (2) Different microcircuit configuration types, identified on the basis of the structural organization of their synapses. (3) Synaptic contacts, not only upon the sensory neurons but also between the efferent fibers themselves. (4) Two identified neurotransmitter candidates, GABA and glutamate. Physiological evidence for GABAergic and glutamatergic transmission is incomplete at spider sensilla. Given that the sensory neurons are quite different in their location and origin, these parallels are most likely convergent. Although their significance is only partially understood, mostly from work on the MRO, the close similarities seem to reflect functional constraints on the organization of efferent pathways in the brain and in the periphery.     

Antennal multiporous sensilla: their gustatory features for host recognition in female parasitic wasps (insecta, Hymenoptera: Platygastroidea).

In parasitic wasps, various kinds of antennal plates have been interpreted as olfactory organs due to the presence of numerous pores. However, on the basis of ultrastructural investigations, some of these multiporous plates were revealed as being release sites of exocrine glands while others were postulated to have a gustatory function. Such sensilla, present only on female antennae, show unique features, found exclusively in Platygastroidea, with some differences between Scelionidae and Platygastridae. The cuticular apparatus consists of a short basiconic shaft with an apical multiporous area, the pores of which are covered by movable structures. The cellular components are made up of a remarkable number of sensory neurons, from 200 to 220, and an undefined number of sheath cells. These multiporous sensilla have tubular accessory glands that release their secretion through the socket sensillum only in scelionids. These morphological characters, combined with behavioral observations, strongly suggest a gustatory function, although electrophysiological studies are still needed to confirm this. A new terminology for antennal multiporous plates is discussed in relation to their different functions.     

Ultrastructure and distribution of antennal sensilla of the chilli thrips Scirtothrips dorsalis hood (Thysanoptera: Thripidae)

The chilli thrips, Scirtothrips dorsalis Hood, is a serious pest of numerous important vegetable and ornamental crops. Various signals, especially phytochemical cues, determine the behavior of the phytophagous thrips at host selection. The sensory abilities of S. dorsalis are poorly understood although the antennae of adult are known to possess important sensory structures in orther insects. In this study, the morphology, distribution, and ultrastructure of the antennal sensilla of the S. dorsalis were examined by using scanning and transmission electron microscopy. Microscopy observations revealed that adult male and female S. dorsalis possess filiform antennae. Each antenna comprises a scape, a pedicel, and a flagellum composed of six segments without clear sexual dimorphism in the number and distribution of antennal sensilla. The scape and pedicel exhibit Böhm's bristles, sensilla chaetica, and sensilla campaniform. The external structures of these organs reveal their mechanosensory function. In the flagellum, the most represented sensilla are the multiporous sensilla basiconica, which can be divided into three types of single‐walled olfactory sensilla; three types of sensilla chaetica with mechanosensory and gustatory functions; sensilla coeloconica, which possess hollow cuticular spoke channels and represent double‐walled olfactory sensilla; sensilla capitula and sensilla cavity with thermo‐hygrosensory functions; and aporous sensilla trichodea with smooth cuticula and mechanosensory function. The putative function of described sensilla is discussed in ralation to host plant selection behavior of S. dorsalis.     

Antennal sensilla and ovipositor morphology of the European birch sawfly Arge pullata Zadd (Hymenoptera: Tenthredinidae,Argidae)

Arge pullata Zadd is an important phytophagous pest that damages red birch Betula albo‐sinensis in Hubei Province, South China. Massive ecological and economic losses have been caused by this species, which threatens the ecological security of the Shennongjia Nature Reserve. To investigate the mechanoreception, chemoreception, and oviposition processes of A. pullata, scanning electron microscopy and optical confocal microscopy were used to reveal the typology, morphology, and distribution of ovipositor and antennal sensilla. The results show that A. pullata has clavate antennae and eight types of sensilla in total, including sensilla chaetica, sensilla trichodea (types 1–3), sensilla basiconica, sensilla coeloconica (types 1 and 2), and Böhm's bristles. Sensilla trichodea type 1 distributed only on male antennae; the densities of sensilla trichodea type 2 and sensilla basiconica differed between the sexes. The binding pattern of ovipositor valvulae was discovered, and one type of sensilla chaetica, two types of sensory pits, and tooth‐like cones as well as two types of microtrichia were found in the ovipositor. Based on morphological evidence and research on Hymenoptera, putative functions are suggested to increase our understanding of the mechanisms by which this species finds hosts and mates, and how oviposition takes place. Microsc. Res. Tech. 77:401–409, 2014. © 2014 Wiley Periodicals, Inc.     

Morphology and distribution of antennal sensilla in egg parasitoid,Ooencyrtus nezarae (Hymenoptera: Encyrtidae)

Morphology of antennal sensilla and their distribution were investigated in adults of Ooencyrtus nezarae, an egg parasitoid of Riptortus pedestris, using scanning electron microscopy. Male antennae was found to be significantly greater in overall length than female antennae. The antenna of O. nezarae was composed of the radicula, scape, pedicel, funicle and clava in both sexes, with seven types of sensilla identified: sensillum trichodea; s. finger‐like; s. placoidea; s. chaetica; s. basiconica; s. coeloconica, and s. campaniform. They occur in varying number and distribution along the antennae. Two sensillum types were further categorized into additional subtypes, with two subtypes in s. trichodea and three in s. chaetica. Among all characterized sensilla, s. trichodea subtype 1 and s. placoidea were multiporous, indicating that the primary function of these sensilla is olfactory. Sensillum trichodea was the most abundant sensillum type on the antennae of both sexes. Sexual dimorphism was only observed from the subtype 1 sensilla of s. trichodea in males and the subtype 3 sensilla of s. chaetica in females. The morphological information established in our study may provide useful information for further investigations in sensory physiological function of each morphological type of sensilla and their related behavior in this egg parasitoid.     

Morphological characterization and distribution of antennal sensilla of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) using scanning electron microscopy

Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) is a serious polyphagous pest of various field and horticultural crops. A complete knowledge on the morphological features of antennal sensory structures is essential for efficient semiochemical-based control methods. The external structure and distribution of antennal sensilla in male and female adults of H. armigera were investigated using scanning electron microscopy. Eight distinct morphological types of sensilla were identified in both sexes: sensilla trichodea, sensilla basiconica, sensilla auricillica, sensilla coeloconica (multiporous), sensilla chaetica (uniporous), sensilla styloconica, sensilla squamiformia, and Böhm sensilla (aporous) in varying numbers and distribution along the length of the antennae. Of these sensilla, the most widespread are sensilla trichodea and sensilla basiconica on the antennae of both sexes. Female antennae have comparatively greater number of sensilla trichodea than male antennae. Among eight types of sensilla, sensilla basiconica, auricillica, styloconica type II, squamiformia, and Böhm sensilla were identified and reported for the first time in H. armigera. Sexual dimorphism in H. armigera was mainly detected as the variations in sensilla shape, numbers, and distribution of each type of sensilla. The sexual difference was observed in the numbers of sensilla coeloconica, chaetica, styloconica, and squamiformia per flagellomere. The possible functions of these sensilla were discussed in view of previously reported lepidopteran insects. The findings provide fundamental information on the morphology and distribution of antennal sensory structures in H. armigera. It would be useful for further detailed studies on physiological and behavioral function of each sensillum type and helpful for formulating related pest control methods.     

Cryoimmobilization and three-dimensional visualization of C. elegans ultrastructure

Caenorhabditis elegans is one of the most important genetic systems used in current biological research. Increasingly, these genetics‐based research projects are including ultrastructural analyses in their attempts to understand the molecular basis for cell function. Here, we present and review state‐of‐the‐art methods for both ultrastructural analysis and immunogold localization in C. elegans. For the initial cryofixation, high‐pressure freezing is the method of choice, and in this article we describe two different strategies to prepare these nematode worms for rapid freezing. The first method takes advantage of transparent, porous cellulose capillary tubes to contain the worms, and the second packs the worms in E. coli and/or yeast paste prior to freezing. The latter method facilitates embedding of C. elegans in a thin layer of resin so individual worms can be staged, selected and precisely orientated for serial sectioning followed by immunolabelling or electron tomography.     

Arthropod sensilla: morphology and phylogenetic considerations

The structures of different types of arthropod sensilla are compared and theories regarding the evolution of these sensory organs are presented. Arthropod sensilla are built according to a common plan, and are probably homologous to scolopidia. Certain similarities in the structure of sensilla in different arthropod groups can be the result of adaptations to specific environments. The structure of sensilla in insect groups, which are regarded to be ancestral, do not appear to be less sophisticated than in groups considered to be more advanced. The different types of pore systems, as well as the structural differentiations of insect olfactory sensillar types remain unexplained. Olfactory sensilla display a large degree of similarity among terrestrial arthropods, whereas crustacean sensilla diverge in structure. In holometabolous insects larval sensilla appear to be structurally quite advanced, and more complex than in the adult. During the ontogeny of both sensilla and scolopidia, these are differentiated in an epithelial layer, resulting in the formation of both sensory and enveloping cells. The developmental patterns of sensilla in the studied insect groups are similar. During the development of sensilla apoptotic process are usually active.     

Antennal sensory structures in Scaphoideus titanus Ball (Hemiptera: Cicadellidae)

Scaphoideus titanus Ball (Hemiptera: Cicadomorpha) is a leafhopper vector of a phytoplasma disease, the Flavescence dorée (FD), belonging to the vine yellows group. A scanning and transmission electron microscope study has been carried out to investigate the antennal sensory structures. The first two segments, the scape and the pedicel, are short, covered by cuticular scales and devoid of sensilla, with exception of some scattered hairs on the pedicel. The flagellum consists of a unique, elongated segment in which numerous subunits can be recognized, being separated by a sort of cuticular crown. The proximal five subunits bear most of the sensilla. We discovered the presence of single- and double-walled coeloconic sensilla, campaniform sensilla, basiconic sensilla, and trichoid sensilla. A scolopidium is located within the proximal region of the flagellum. Ultrastructural investigations suggest that the antennal sensilla could be involved in the perception of air-borne vibrations, temperature, and humidity variations. The most relevant feature is the extreme reduction of the olfactory sensilla, both in terms of number of sensory structures and sensory neurons per sensillum. The strong reduction in antennal olfactory sensilla to which this specie has undergone is discussed as possible consequence of the specificity toward the host plant.