Ultrastructural Analysis of the Sensilla of Austroperipatus aequabilis Reid, 1996 (Onychophora,Peripatopsidae)

The head of Austroperipatus aequabilis bears five types of sensilla. which were examined by electron microscopy. They differ from each other in position, shape of outer sensory elements and cuticular socket structures. Thus, we distinguish sensilla with sensory hairs, sensilla with sensory bulbs, cone-shaped sensilla. sensilla with sensory bristles, and sensilla of the lips. They are composed of up to 15 cells, which can he separated into four cell types. The most frequent cell type is the bipolar receptor cell that occurs in all sensilla. The apical surface of this primary receptor cell is characterized by one or two partly branched cilia with a basal 9 × 2 + 0 pattern of microtubules. A modified bipolar receptor cell was found in all sensilla bearing a sensory peg except for the sensilla equipped with sensory bristles. The apical dendrite extends to a long pale process which exclusively contains mitochondria and single microtubules. In all sensilla examined in this study at least one supporting cell occurs which is characterized by parallel microvilli. An additional function of this cell type as a part of the stimulus-conducting system is possible. In the sensillum with a sensory bulb two kinds of supporting cells occur. A unique cell type with an upside down position has regularly been found in all sensilla bearing a sensory peg. Apart from the sensilla they also occur within the labial epidermis. Since most sensilla contain several different receptor cells, they can be considered as complex sense organs. © 1998 The Royal Swedish Academy of Sciences. Published by Elsevier Science Ltd. All rights reserved     

Fine structure of the sensilla of Peripatopsis moseleyi (Onychophora)

Summary Three types of sensilla occurring on the lips and on the antennae of Peripatopsis moseleyi have been investigated by scanning and transmission electron microscopy. On the lips sensory spines can be found which contain numerous cilia originating from bipolar receptor cells. They reach the tip of the spine where the cuticle is modified. The perikarya of the sensory cells, a large supporting cell with a complicated surface and a second type of receptor, form a bud-like structure and are surrounded by a layer of collagen fibrils. The second receptor cell bears apical stereocilia as well as a kinocilium which are directed towards the centre of the animal — thus the cell appears to be turned upside down. The sensilla of the antennae are 1) sensory bristles containing two or three kinds of receptor cells, one of which bears an apical cilium and one kind of supportive cell and 2) sensory bulbs located within furrows consisting of receptor cells with branched cilia and two kinds of supportive cells which are covered by a modified thin cuticle. According to the electron microscopical findings the sensory spines on the lips are presumably chemoreceptors. The sensory bristles on the antennae can be regarded as mechanoreceptors and the sensory bulbs as chemoreceptors.Supported by the Deutsche Forschungsgemeinschaft (Sto 75/3)     

The External Structure and Distribution of Sensilla in the Medicinal Leech

Abstract The body of Hirudo medicinalis consists of 32 segments. The quinquannulate midbody segments 3–18 bear 14 sensilla on the central (neural) annulus. Elsewhere segments are represented by fewer than five annuli but sensilla are retained on the neural annulus. From neural sensilla protrude S hairs (cilia from uniciliate cells) which are thought to detect water currents. Two categories of multiciliate cell, of unknown function, are also present within neural sensilla; (i) grouped cilia extending beyond the cuticle (G hairs), and (ii) cilia which radiate out beneath the cuticle. Studies of the entire external surface of leech body wall with a scanning electron microscope revealed the presence of large numbers of sensilla on every annulus including the neural annuli. These sensilla lack S hairs. Our results show: (i) annular sensilla have a significantly smaller surface area than neural sensilla (p < 0.001 Mann-Whitney test, 2 tailed), (ii) The position and number of the small sensilla varies from annulus to annulus, segment to segment and individual to individual, (iii) Significantly higher numbers of small sensilla were found where non-neural annuli remain single and have not undergone a further division during development as happens in the quinquannulate segments. The data suggest that small sensilla continue to be added during the adult life of the leech.     

Cytoskeletal elements in arthropod sensilla and mammalian photoreceptors.

Ciliary receptor cells, typified by cilia or modified cilia, are very common in the animal kingdom. In addition to the cytoskeleton of their ciliary processes these receptors possess other specific prominent cytoskeletal elements. Two representative systems are presented: i) scolopidia, mechanosensitive sensilla of various arthropod species; and ii) photoreceptor cells of the retina of the bovine eye. Two cytoskeletal structures are characteristic for arthropod scolopidia: a scolopale typifies the innermost auxiliary cell, and long ciliary rootlets are extending well into the sensory cells. The latter element is also characteristic for the inner segment of the photoreceptor cells in bovine. The scolopale of scolopidia is mainly composed of actin filaments. In the absence of myosin, the uniform polarity of the actin filaments and their association with tropomyosin all indicate a stabilizing role of the filament bundles within the scolopale. This function and a certain elasticity of actin filament bundles may be important during stimulation of the sensilla. The ciliary rootlets of both systems originate at the basal bodies at the ciliary base of the sensory cells and project proximally. These rootlets are composed of longitudinally oriented, fine filaments forming a characteristic regular cross-striation. An alpha-actinin immunoreactivity was detected within the ciliary rootlets of scolopidia. In addition, antibodies to centrin react with the rootlets of both types of receptors. Since centrin is largely responsible for the contraction of the flagellar rootlets in green algae, contraction may also occur in the ciliary rootlets of insect sensilla and vertebrate photoreceptors. In both systems, contraction or relaxation of the ciliary rootlets could serve in sensory transduction or adaptation.     

Morphology of chemosensory organs required for feeding in the leech Hirudo medicinalis

Sensilla that line the upper edge of the lip in the leech Hirudo medicinalis and that contain chemoreceptors required for feeding were examined in the scanning and transmission electron microscopes. The sensilla include two size-classes of ciliated button-like mounds--one about 35 microns in diameter and another about 10 microns in diameter. The larger sensilla are at the center of unpigmented patches of skin which are visible in the light microscope, while the smaller sensilla have not been previously described as distinct structures. Electron microscopy, though not light microscopy, shows that the lip sensilla differ markedly from the segmental sensilla of the leech, which have been shown to mediate mechanoreception and photoreception. In particular, the chemosensory lip sensilla contain multiciliated cells with cilia of a uniform length, whereas the segmental sensilla contain uniciliated cells with long, whip-like cilia, as well as multiciliated cells with short, stiff cilia. Thus, the two types of sensilla differ morphologically as well as functionally. In addition to the ciliated sensilla along the upper lip, structures consisting of a short, club-like process surrounded by granular material were observed inside the mouth. These structures may also be chemosensory organs.     

Structure and function of vertebrate cilia, towards a new taxonomy

In this review, we propose a new classification of vertebrate cilia/flagella and discuss the evolution and prototype of cilia. Cilia/flagella are evolutionarily well-conserved membranous organelles in eukaryotes and serve a variety of functions, including motility and sensation. Vertebrate cilia have been traditionally classified into conventional motile cilia and sensory primary cilia. However, an avalanche of emerging evidence on the variations of cilia has made it almost impossible to classify them in a simple dichotomic manner. For example, conventional motile cilia are also involved in the sensation of bitter taste to facilitate the beating of cilia as a defense system of the respiratory system. On the other hand, the primary cilium, often regarded as a non-motile sensory organelle, has been revealed to be motile in vertebrate embryonic nodes, where they play a crucial role in the determination of left-right asymmetry of the body. Moreover, choroid plexus epithelial cells in the cerebral ventricular system exhibit multiple primary cilia on a single cell. Considering these lines of evidence on the diversity of cilia, we believe the classification of cilia should be based on their structure and function, and include more detailed criteria. Another intriguing issue is how in the evolution of cilia, their function and morphology are combined. For example, has motility been acquired from originally sensory cilia, or vice versa? Alternatively, were they originally hybrid in nature? These questions are inseparable from the classification of cilia per se. We would like to address these conundrums in this review article, principally from the standpoint of differentiation of the animal cell.     

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。     

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.     

Ciliated sensory receptors of the unactivated metacestode of Hymenolepis microstoma

The fine structure of the ciliated sensory endings of the unactivated metacestode of Hymenolepis microstoma is described. The type I sensilla, localized in areas of the scolex and sucker tegument, are characterized by a long cilium (1.20 × 0.20 μ) embedded in a bulb containing two electron-dense collars and three to five mitochondria. The type II sensilla, located in tegumentary pits of the rostellum, are characterized by a short cilium (0.62 × 0.28 μ) possessing ciliary rootlets, embedded in a bulb containing a single electron-dense collar and one to three mitochondria. Both sensory cilia are characterized by a 9 + 6 + 1 microtubular substructure. The sensory bulbs contain large numbers of membrane bound electron-lucent vesicles; neurotubules are absent at this stage of development.     

Antennal sensory organs in the millipede Eurypauropus ornatus: Fine structure of the flagella and globulus

On the antennal tip of Eurypauropus ornatus are 3 threadlike sensilla—the flagella, and a single spheroid sensillum—the globulus. Each of the 3 flagella is innervated by 2 groups of sensory cells. One group contains 4 cells, the other, 5. All cells of the “four group” and 3 of the “five group” are comprised of single cilia and unbranched dendrites which extend along the lumen of the flagellum. Two cells of the “five group” have double cilia and pairs of unbranched dendrites. One pair also enters the flagellum and the other pair terminates beneath the flagellar base to form a concentric array of lamellae. No pores are present in the cuticular wall. Eight sensory cells innervate the globulus. They are arranged in 3 groups, one triplet and 2 pairs, in addition to a single cell. The single cell contains a pair of cilia whose unbranched dendrites differentiate into tubular bodies that are inserted into the base of the globulus. Each of the other 7 sensory cells has a single cilium. Their unbranched dendrites penetrate into the globulus in 3 groups as described for the sensory cells. The dendrites in each group terminate in an individual pore channel at the globulus tip and completely fuse with the electron-dense material that plugs the pore channel. Based on structural similarities to sensilla having known functions, it is probable that the flagella and the globulus are chemoreceptors, the former responding to odors, the latter sensitive to substances in aqueous solution.     

Ultrastructural investigation of the nuchal organs ofPygospio elegans (Polychaeta). I. Larval nuchal organs

The structural differentiation of the nuchal organs during the post-embryonic development ofPygospio elegans is described. The sensory organs are composed of two cell types: ciliated cells and bipolar primary sensory cells, constituting the nuchal ganglion, which is associated with both the sensory epithelium and the brain. Since the sensory neurons are largely integrated into posterolateral parts of the cerebral ganglion, the nuchal organs are primary presegmental structures. The microvilli of the ciliated cells form a cover over the cuticle with a presumed protective function. An extracellular space extends between cuticle and sensory epithelium. The distal dendrites of the sensory cells terminate in sensory bulbs, bearing one modified sensory cilium each that projects into the olfactory chamber, embedded within the secretion of the ciliated cells. During development, the nuchal organs increase in size. This is accompanied by a shift in position, an expansion of the sensory area, and secretory activity of the ciliated cells. The nuchal ganglion differentiates into three nuchal centres forming three distinct sensory areas around the ciliated region. Each nuchal complex reveals two short nuchal nerves comprising the sensory axons, which enter the posterior circumesophageal connective. The sensory cells lying in the brain exhibit neurosecretory activity; the sensory cilia enlarge their surface area by dilating and branching. Nuchal organs accomplish the basic structural adaptions of chemoreceptors and show structural analogies to arthropod olfactory sensilla; thus, there is every reason to suppose chemoreceptor function.     

The coelocapitular sensillum,an antennal hygro- and thermoreceptive sensillum of the honey bee,Apis mellifera L.

Summary The sensillum coelocapitulum, a hygro- and thermoreceptive sensillum of the honey bee, Apis mellifera, was investigated by electron microscopy. The cuticular apparatus of the sensillum is a mushroomshaped protrusion, devoid of pores, set in a narrow cylindrical pit positioned centrally within a cuticular, shallow depression. There may be three or four receptor cells. Three receptor cells have unbranched sensory cilia, containing densely packed microtubules, which extend distally into the cuticular apparatus and completely fill its cavity. These connecting cilia are of the usual 9+0 type. The fourth receptor, if present, has a thin sensory cilium which terminates beneath the cuticular apparatus. Its connecting cilium has armed outer doublets. The outer cavity is formed by two enveloping cells and is completely sealed off. Lipid deposits are present within the cavity and the tormogen cell. The thecogen cell has scolopale rod-like structures around the inner cavity. Features common to the insect hygro- and thermoreceptive sensilla are discussed in comparison with those of other insects.     

Mutant sensory cilia in the nematode Caenorhabditis elegans

Eight classes of chemosensory neurons in C. elegans fill with fluorescein when living animals are placed in a dye solution. Fluorescein enters the neurons through their exposed sensory cilia. Mutations in 14 genes prevent dye uptake and disrupt chemosensory behaviors. Each of these genes affects the ultrastructure of the chemosensory cilia or their accessory cells. In each case, the cilia are shorter or less exposed than normal, suggesting that dye contact is the principal factor under selection. Ten genes affect many or all of the sensory cilia in the head. The daf-19 (m86) mutation eliminates all cilia, leaving only occasional centrioles in the dendrites. The cilia in che-13 (e1805), osm-1 (p808), osm-5 (p813), and osm-6 (p811) mutants have normal transition zones and severely shortened axonemes. Doublet-microtubules, attached to the membrane by Y links, assemble ectopically proximal to the cilia in these mutants. The amphid cilia in che-11 (e1810) are irregular in diameter and contain dark ground material in the middle of the axonemes. Certain mechanocilia are also affected. The amphid cilia in che-10 (e1809) apparently degenerate, leaving dendrites with bulb-shaped endings filled with dark ground material. The mechanocilia lack striated rootlets. Cilia defects have also been found in che-2, che-3, and daf-10 mutants. The osm-3 (p802) mutation specifically eliminates the distal segment of the amphid cilia. Mutations in three genes affect sensillar support cells. The che-12 (e1812) mutation eliminates matrix material normally secreted by the amphid sheath cell. The che-14 (e1960) mutation disrupts the joining of the amphid sheath and socket cells to form the receptor channel. A similar defect has been observed in daf-6 mutants. Four additional genes affect specific classes of ciliated sensory neurons. The mec-1 and mec-8 (e398) mutations disrupt the fasciculation of the amphid cilia. The cat-6 (e1861) mutation disrupts the tubular bodies of the CEP mechanocilia. A cryophilic thermotaxis mutant, ttx-1 (p767), lacks fingers on the AFD dendrite, suggesting this neuron is thermosensory.     

The Nervous System of the Male Dinophilus gyrociliatus (Annelida: Polychaeta). I. Number, Types and Distribution Pattern of Sensory Cells

The entire nervous system of the smallest annelid hitherto known, the dwarf male of the highly dimorphic species Dinophilus gyrociliatus , has been reconstructed by means of TEM investigations of serial ultrathin sections. Altogether there are 68 neurons, 40 of which have a sensory function. The structure and distribution of them is described. The receptor endings of the 20 sensory cells of each side are located either in two groups — the anterior receptor group and the posterior receptor group — or are singly positioned in the integument. Structural differences of the apical portion of the dendrites enables four types of receptors to be distinguished: three types with emergent cilia and one type with non-emergent cilia. Neurons with emergent cilia can be monociliated collar cells as well as mono- or multiciliated cells without collar. Special vesicle-in-vesicle structures, are located close to the basal portion of the cilia in some of these cells. The non-emergent cilia border closely to a neighbouring epidermal cell and contain a prominent intraciliary vesicle. The function of receptors is discussed with regard to a comparison with receptors in other polychaete species, structural specializations and their distribution pattern on the animal's surface.     

Fine Structure of Antennal Sensilla of Paysandisia archon and Electrophysiological Responses to Volatile Compounds Associated with Host Palms

Paysandisia archon (Lepidoptera: Castniidae) is a serious pest of palm trees. A comprehensive knowledge of the insect olfactory system is essential for the development of efficient semiochemical-based control methods. The olfactory sensilla are located particularly on the antennae, and these can detect plant volatiles that provide important cues for the insects in the search for their host plants. To date, the fine structure of P. archon antennal sensilla studies and their role in host-plant perception have not been investigated in great detail. Using light microscopy and scanning and transmission electron microscopy, the antennae of both sexes of P. archon are described here in detail, according to the different types, quantities and distributions of the sensilla. Six types of sensilla were identified. The most widespread are sensilla trichoidea, sensilla basiconica and sensilla auricilica, which are associated with olfactory function. These have cuticular shafts characterised by numerous pores, and they are innervated by two or three sensory neurons. Sensilla coeloconica, sensilla chaetica and sensilla ampullacea are associated with olfactory or olfactory-thermoreception, mechano-gustatory, and thermo-hygroreception functions, respectively. Moreover, the role of P. archon antennae in locating of the host palms was evaluated using electroantennograms, to monitor responses to ester and terpene compounds previously identified as volatiles of damaged/fermenting palm tissues. P. archon showed responses to all of the synthetic chemicals tested, with greater responses in the females, providing a significant sex*dose effect. Among the compounds tested, ethyl isobutyrate elicited the strongest antenna responses. The fine structure of the cuticular and cellular components of the P. archon antenna sensory equipment is described for the first time. The results of this study form an important starting point and complement physiological and behavioural studies, to provide valuable information of practical importance for the development of efficient semiochemical-based control methods.     

雄蚤生殖节的扫描电镜观察

本文利用扫描电镜对雄蚤生殖节表面细微结构进行了观察,发现了一些光镜下不易看到的纹理状结构和小棘及一些感器.这些微小的纹理状结构和小棘在不同蚤种之间有差异,这种差异具有相对的稳定性,可望用于蚤的亚显微水平的分类.通过对感器的观察,发现生殖节上具有大量的毛形感器,另外还观察到有锥形感器、短锥形感器、钟形感器、栓锥感器、腔锥感器及刺形感器等.这些感器的数量分布及有无因种而异.这些感器的存在与交配行为密切相关,作者对此也进行了讨论.     

Distribution and fine structure of antennal olfactory sensilla in Japanese dung beetles,Geotrupes auratus Mtos. (Coleoptera : Geotrupidae) and Copris pecuarius Lew. (Coleoptera : Scarabaeidae)

The distribution and fine structure of olfactory sensilla on the antennal flagella of the Japanese dung beetles, Geotrupes auratus (Coleoptera : Geotrupidae) and Copris pecuarius (Coleoptera : Scarabaeidae) were studied by scanning and transmission electron microscopy. In both types of dung beetles, sensilla on the antennal segments were most abundant on the apical 3 lamellar segments, where sensilla basiconica were particularly concentrated. The sensilla were solely concentrated in the central part of the distal side of both the 9th and 10th segments in G. auratus and the 7th and 8th segments in C. pecuarius, respectively. The numbers of sensilla in these areas (channels) of the 9th and 10th segments in G. auratus were approximately 4,500 and 3,000, respectively. In C. pecuarius, there were about 1,900 sensilla on the 7th segment and l,700 sensilla on the 8th. The cuticles of the sensilla in both species were perforated by numerous elongate pores, with a pore density of about 45/ μm² in G. auratus and 30/ μm² in C. pecuarius. Each sensillum basiconicum in C. pecuarius was generally associated with 2 sensory neurons. The sensory cilia passed into the sensillum lumen and branched profusely, occupying most of its space. On the basis of the fine structure and dense location, these sensilla are considered to have an olfactory function for food detection.     

Tissue expression patterns identify mouse cilia genes.

In mammals, cilia are critical for development, sensation, cell signaling, sperm motility, and fluid movement. Defects in cilia are causes of several congenital syndromes, providing additional reasons to identify cilia-related genes. We hypothesized that mRNAs selectively abundant in tissues rich in highly ciliated cells encode cilia proteins. Selective abundance in olfactory epithelium, testes, vomeronasal organ, trachea, and lung proved to be an expression pattern uniquely effective in identifying documented cilia-related genes. Known and suspected cilia-related genes were statistically overrepresented among the 99 genes identified, but the majority encoded proteins of unknown function, thereby predicting new cilia-related proteins. Evidence of expression in a highly ciliated cell, the olfactory sensory neuron, exists for 73 of the genes. In situ hybridization for 17 mRNAs confirmed expression of all 17 in olfactory sensory neurons. Most were also detected in vomeronasal sensory neurons and in neighboring tissues rich in ciliated cells such as respiratory epithelium. Immunoreactivity for one of the proteins identified, Spa17, colocalized with acetylated tubulin in the cilia layer of the olfactory epithelium. In contrast, the ciliary rootlet protein, Crocc, was located in discrete structures whose position was consistent with the dendritic knobs of the olfactory sensory neurons. A compilation of >2,000 mouse genes predicted to encode cilia-related proteins revealed a strong correlation (R = 0.99) between the number of studies predicting a gene's involvement in cilia and documented evidence of such involvement, a fact that simplifies the selection of genes for further study of the physiology of cilia.