Neurogenic and antineurogenic effects from modifications at the Notch locus

The best studied mutations at the Notch locus produce a neurogenic phenotype, with a massive overgrowth of the nervous system at the expense of epidermis. We report here that, in the development of the adult peripheral nervous system, the Abruptex alleles of Notch have the opposite phenotype, namely an underproduction of sensory organs or sensilla. This arises primarily not from an arrest of the lineages that produce sensilla, from the degeneration of sensillar cells, or from the transformation into neurons of cells that normally secrete the cuticular components of a sensillum (as can happen in Notch alleles). Rather, our evidence argues strongly that the sensillar mother cells never form. This implies that the Notch protein plays a role in the process that first generates a difference between sensillar mother cells and ordinary epidermal cells. The number of sensilla formed on the wing of flies carrying multiple doses of Notch+ is virtually the same as that of wild type, i.e. the Abruptex phenotype is not reproduced to any significant extent. This suggests that the single amino acid substitutions that occur in Abruptex mutants confer on the protein some functionally distinctive feature, possibly more powerful intermolecular binding or altered stability.     

Echinoid mutants exhibit neurogenic phenotypes and show synergistic interactions with the Notch signaling pathway

During neurogenesis in Drosophila, groups of ectodermal cells are endowed with the capacity to become neuronal precursors. The Notch signaling pathway is required to limit the neuronal potential to a single cell within each group. Loss of genes of the Notch signaling pathway results in a neurogenic phenotype: hyperplasia of the nervous system accompanied by a parallel loss of epidermis. Echinoid (Ed), a cell membrane associated Immunoglobulin C2-type protein, has previously been shown to be a negative regulator of the EGFR pathway during eye and wing vein development. Using in situ hybridization and antibody staining of whole-mount embryos, we show that Ed has a dynamic expression pattern during embryogenesis. Embryonic lethal alleles of ed reveal a role of Ed in restricting neurogenic potential during embryonic neurogenesis, and result in a phenotype similar to that of loss-of-function mutations of Notch signaling pathway genes. In this process Ed interacts closely with the Notch signaling pathway. Loss of ed suppresses the loss of neuronal elements caused by ectopic activation of the Notch signaling pathway. Using a temperature-sensitive allele of ed we show, furthermore, that Ed is required to suppress sensory bristles and for proper wing vein specification during adult development. In these processes also, ed acts in close concert with genes of the Notch signaling pathway. Thus the extra wing vein phenotype of ed is enhanced upon reduction of Delta (Dl) or Enhancer of split [E(spl)] proteins. Overexpression of the membrane-tethered extracellular region of Ed results in a dominant-negative phenotype. This phenotype is suppressed by overexpression of E(spl)m7 and enhanced by overexpression of Dl. Our work establishes a role of Ed during embryonic nervous system development, as well as adult sensory bristle specification and shows that Ed interacts synergistically with the Notch signaling pathway.     

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.     

黑胸大蠊下颚须和下唇须上感器扫描电镜观察

利用扫描电子显微镜观察了黑胸大蠊Periplanetafuliginosa(Serville)成虫下颚须和下唇须上的感器形态。结果发现,在黑胸大蠊下颚须和下庸须末节顶端何感器密集区,尤其是下颚须第5节内侧顶端,有一狭长沟壑,内有大量的带槽锥形感器。通过重点观察感器密集区,发现主要有5~6种类型感器,分别为带槽锥形、毛形、刺形、钟形、齿状、针形感器,其中有些感器又可分为几种亚类型。比较研究发现下颚须和下唇须上感器类型除了带槽锥形感器以外基本相似,只是数量上有区别。     

The Drosophila IgC2 domain protein Friend-of-Echinoid,a paralogue of Echinoid,limits the number of sensory organ precursors in the wing disc and interacts with the Notch signaling pathway

The Notch signaling pathway is critical in cell fate specification throughout development. In the developing wing disc, single sensory organ precursors (SOPs) are selected from proneural clusters via a process of lateral inhibition mediated by the Notch signaling pathway. The epidermal growth factor receptor (EGFR) pathway has also been implicated in SOP formation. Here, we describe the Drosophila melanogaster gene friend of echinoid (fred), a paralogue of echinoid (ed), a gene recently identified as a negative regulator of the EGFR pathway. fred function was examined in transgenic flies by using inducible RNA interference (RNAi). Suppression of fred in developing wing discs results in specification of ectopic SOPs, additional microchaeta, and cell death. In eye-antennal discs, fred suppression causes a rough eye phenotype. These phenotypes are suppressed by overexpression of Notch, Suppressor of Hairless [Su(H)], and Enhancer of split m7. In contrast, overexpression of Hairless, a negative regulator of the Notch pathway, and decreased Su(H) activity enhance these phenotypes. Thus, fred acts in close concert with the Notch signaling pathway. Dosage-sensitive genetic interaction also suggests a close relationship between fred and ed.     

Adaptation of sensory systems of gamasid mites (Parasitiformes, Gamasina) to dwelling in different environment

The main complication sensory organs (the palpal organ and the tarsal sensory complex) of several species of gamasid mites were studied in scanning electron microscope. The species examined included permanent ectoparasites (Laelaps agilis, Laelaptidae), parasites of the nasal cavity and respiratory tract of birds (Sternostoma tracheocolum and Ptilonyssus reguli, Rhinonyssidae), dwellers of the sea littoral zone (Parasitus kempersi, Parasitus immanis, Parasitidae), and mites found on soil and on plants (Amblyseius barkeri, Parasitidae). Similar sensillar types, including olfactory SW-WP sensilla, contact chemo-mechanosensory (SW-UP and DW-UP) sensilla, termo-chemo-mechanosensitive (DW-WP) sensilla of two types, and tactile (NP) sensilla were found in all these species, excluding endoparasites, where some sensillar types (in particular, DW-WP sensilla with slit-like pores) are absent. It was shown that the topography of olfactory SW-WP sensilla of the tarsal complex reflects taxonomic position and phylogenetic history of mite genera, whereas the number of certain sensillar types and the degree of their development reflect ecological specialization of species. The palpal organ is characterized by rather uniform structure in mites of different families, dwellers of different environments, except for the endoparasites of the family Rhinonyssidae, where this organ is strongly reduced.     

The development of the sensory neuron pattern in the antennal disc of wild-type and mutant (lz3, ssa) Drosophila melanogaster

The development of the sensory neuron pattern in the antennal disc of Drosophila melanogaster was studied with a neuron-specific monoclonal antibody (22C10). In the wild type, the earliest neurons become visible 3 h after pupariation, much later than in other imaginal discs. They lie in the center of the disc and correspond to the neurons of the adult aristal sensillum. Their axons join the larval antennal nerve and seem to establish the first connection towards the brain. Later on, three clusters of neurons appear in the periphery of the disc. Two of them most likely give rise to the Johnston's organ in the second antennal segment. Neurons of the olfactory third antennal segment are formed only after eversion of the antennal disc (clusters t1-t3). The adult pattern of antennal neurons is established at about 27% of metamorphosis. In the mutant lozenge3 (lz3), which lacks basiconic antennal sensilla, cluster t3 fails to develop. This indicates that, in the wild type, a homogeneous group of basiconic sensilla is formed by cluster t3. The possible role of the lozenge gene in sensillar determination is discussed. The homeotic mutant spineless-aristapedia (ssa) transforms the arista into a leg-like tarsus. Unlike leg discs, neurons are missing in the larval antennal disc of ssa. However, the first neurons differentiate earlier than in normal antennal discs. Despite these changes, the pattern of afferents in the ectopic tarsus appears leg specific, whereas in the non-transformed antennal segments a normal antennal pattern is formed. This suggests that neither larval leg neurons nor early aristal neurons are essential for the outgrowth of subsequent afferents.     

The Sno Oncogene Antagonizes Wingless Signaling during Wing Development in Drosophila

The Sno oncogene (Snoo or dSno in Drosophila) is a highly conserved protein and a well-established antagonist of Transforming Growth Factor-β signaling in overexpression assays. However, analyses of Sno mutants in flies and mice have proven enigmatic in revealing developmental roles for Sno proteins. Thus, to identify developmental roles for dSno we first reconciled conflicting data on the lethality of dSno mutations. Then we conducted analyses of wing development in dSno loss of function genotypes. These studies revealed ectopic margin bristles and ectopic campaniform sensilla in the anterior compartment of the wing blade suggesting that dSno functions to antagonize Wingless (Wg) signaling. A subsequent series of gain of function analyses yielded the opposite phenotype (loss of bristles and sensilla) and further suggested that dSno antagonizes Wg signal transduction in target cells. To date Sno family proteins have not been reported to influence the Wg pathway during development in any species. Overall our data suggest that dSno functions as a tissue-specific component of the Wg signaling pathway with modest antagonistic activity under normal conditions but capable of blocking significant levels of extraneous Wg, a role that may be conserved in vertebrates.     

Functional anatomy of sensory structures on the antennae of Psylliodes chrysocephala L. (Coleoptera: Chrysomelidae)

The ultrastructure and distribution of sensilla on the antennae of the cabbage stem flea beetle, Psylliodes chrysocephala, were investigated using scanning and transmission electron microscopy techniques. Eight different sensillar types were distinguished. These were; hair plate sensilla, sensilla chaetica, three types of sensilla trichodea, sensilla basiconica, grooved peg sensilla and styloconic sensilla. The sensilla chaetica are known to be gustatory receptors. Ultrastructure indicates that the hair plate sensilla and sensilla trichodea type one are probably mechanoreceptors, whilst the sensilla styloconica are probably thermo-hygro receptors. These thermo-hygroreceptors are unusual in that they are innervated by two sensory cells (one hygroreceptor and one thermoreceptor) rather than the more usual triad. The remaining four sensillar types all have a porous hair shaft, indicating an olfactory role. One of these (the grooved peg sensillum) may also have a thermoreceptive function. No sexual dimorphism was found in the structure, number or distribution of the antennal sensilla.     

Physiology and glomerular projections of olfactory receptor neurons on the antenna of female Heliothis virescens (Lepidoptera: Noctuidae) responsive to behaviorally relevant odors

The neurophysiology and antennal lobe projections of olfactory receptor neurons housed within short trichoid sensilla of female Heliothis virescens F. (Noctuidae: Lepidoptera) were investigated using a combination of cut-sensillum recording and cobalt-lysine staining techniques. Behaviorally relevant odorants, including intra- and inter-sexual pheromonal compounds, plant and floral volatiles were selected for testing sensillar responses. A total of 184 sensilla were categorized into 25 possible sensillar types based on odor responses and sensitivity. Sensilla exhibited both narrow (responding to few odors) and broad (responding to many odors) response spectra. Sixty-six percent of the sensilla identified were stimulated by conspecific odors; in particular, major components of the male H. virescens hairpencil pheromone (hexadecanyl acetate and octadecanyl acetate) and a minor component of the female sex pheromone, (Z)-9-tetradecenal. Following characterization of the responses, olfactory receptor neurons within individual sensilla were stained with cobalt lysine (N=39) and traced to individual glomeruli in the antennal lobe. Olfactory receptor neurons with specific responses to (Z)-9-tetradecenal, a female H. virescens sex pheromone component, projected to the female-specific central large female glomerulus (cLFG) and other glomeruli. Terminal arborizations from sensillar types containing olfactory receptor neurons sensitive to male hairpencil components and plant volatiles were also localized to distinct glomerular locations. This information provides insight into the representation of behaviorally relevant odorants in the female moth olfactory system. Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Tufted is a gain-of-function allele that promotes ectopic expression of the proneural gene amos in Drosophila

The Tufted(1) (Tft(1)) dominant mutation promotes the generation of ectopic bristles (macrochaetae) in the dorsal mesothorax of Drosophila. Here we show that Tft(1) corresponds to a gain-of-function allele of the proneural gene amos that is associated with a chromosomal aberration at 36F-37A. This causes ectopic expression of amos in large domains of the lateral-dorsal embryonic ectoderm, which results in supernumerary neurons of the PNS, and in the notum region of the third instar imaginal wing, which gives rise to the mesothoracic extra bristles. Revertants of Tft(1), which lack ectopic neurons and bristles, do not show ectopic expression of amos. One revertant is a loss-of-function allele of amos and has a recessive phenotype in the embryonic PNS. Our results suggest that both normal and ectopic Tft(1) bristles are generated following similar rules, and both are subjected to Notch-mediated lateral inhibition. The ability of Tft(1) bristles to appear close together may be due to amos having a stronger proneural capacity than that of other proneural genes like asense and scute. This ability might be related to the wild-type function of amos in promoting development of large clusters of closely spaced olfactory sensilla.     

Functionally redundant peg sensilla on the scorpion pecten

All scorpions have two mid-ventral organs called pectines. Each pecten has thousands of pore-tipped sensilla sensitive to a variety of volatile organic and water-based stimulants. However, it was previously unknown whether individual sensilla were functionally identical or different. The information enhancement hypothesis predicts that all sensilla have similar chemosensitivities such that each is a unit of a parallel processing system. The information segmentation hypothesis states that sensilla differ in their chemosensitivities, a functional arrangement akin to the glomeruli-specific chemical detection system in the moth or human olfactory sense. In this study, we tested these hypotheses by extracellularly tip-recording sensillar responses to three aqueous tastants: 0.01 M KCl, 0.1 M citric acid, and 40% ethanol by volume. We isolated stimulation to one sensillum at a time and compared the chemoresponses. Sensilla appeared to respond similarly to the same stimulant (i.e., sensillar tip-recordings revealed activity of the same cell types), although sometimes a few sensilla responded with higher spike rates than the others. We conclude that our data primarily support the information enhancement hypothesis but for future tests of sensillar function we suggest a new hybrid model, which proposes that a few specialized sensilla exist among a mostly uniform field of identical sensilla.     

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

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

The organization of the chemosensory system in Drosophila melanogaster: a rewiew

This review surveys the organization of the olfactory and gustatory systems in the imago and in the larva of Drosophila melanogaster, both at the sensory and the central level. Olfactory epithelia of the adult are located primarily on the third antennal segment (funiculus) and on the maxillary palps. About 200 basiconic (BS), 150 trichoid (TS) and 60 coeloconic sensilla (CS) cover the surface of the funiculus, and an additional 60 BS are located on the maxillary palps. Males possess about 30% more TS but 20% fewer BS than females. All these sensilla are multineuronal; they may be purely olfactory or multimodal with an olfactory component. Antennal and maxillary afferents converge onto approximately 35 glomeruli within the antennal lobe. These projections obey precise rules: individual fibers are glomerulus-specific, and different types of sensilla are associated with particular subsets of glomeruli. Possible functions of antennal glomeruli are discussed. In contrast to olfactory sensilla, gustatory sensilla of the imago are located at many sites, including the labellum, the pharynx, the legs, the wing margin and the female genitalia. Each of these sensory sites has its own central target. Taste sensilla are usually composed of one mechano-and three chemosensory neurons. Individual chemosensory neurons within a sensillum respond to distinct subsets of molecules and project into different central target regions. The chemosensory system of the larva is much simpler and consists essentially of three major sensillar complexes on the cephalic lobe, the dorsal, terminal and ventral organs, and a series of pharyngeal sensilla.     

The organization of the chemosensory system in Drosophila melanogaster: a rewiew

This review surveys the organization of the olfactory and gustatory systems in the imago and in the larva of Drosophila melanogaster, both at the sensory and the central level. Olfactory epithelia of the adult are located primarily on the third antennal segment (funiculus) and on the maxillary palps. About 200 basiconic (BS), 150 trichoid (TS) and 60 coeloconic sensilla (CS) cover the surface of the funiculus, and an additional 60 BS are located on the maxillary palps. Males possess about 30% more TS but 20% fewer BS than females. All these sensilla are multineuronal; they may be purely olfactory or multimodal with an olfactory component. Antennal and maxillary afferents converge onto approximately 35 glomeruli within the antennal lobe. These projections obey precise rules: individual fibers are glomerulus-specific, and different types of sensilla are associated with particular subsets of glomeruli. Possible functions of antennal glomeruli are discussed. In contrast to olfactory sensilla, gustatory sensilla of the imago are located at many sites, including the labellum, the pharynx, the legs, the wing margin and the female genitalia. Each of these sensory sites has its own central target. Taste sensilla are usually composed of one mechano-and three chemosensory neurons. Individual chemosensory neurons within a sensillum respond to distinct subsets of molecules and project into different central target regions. The chemosensory system of the larva is much simpler and consists essentially of three major sensillar complexes on the cephalic lobe, the dorsal, terminal and ventral organs, and a series of pharyngeal sensilla.     

豌豆潜蝇姬小蜂雌蜂触角感器的扫描电镜观察

应用扫描电镜观察豌豆潜蝇姬小蜂Diglyphus isaea Walker雌蜂触角感器,包括Bhm氏鬃毛、感觉孔、单孔毛状感器、多孔片状感器、锥形乳头状感器、厚壁化学感器、薄壁化学感器和毛状感器8种感器,分别对这8种感器的形态特征进行详细描述,并推测其可能的功能;还对有关的潜蝇姬小蜂生物学特性做深入的分析和讨论。     

Comparative external ultrastructure and diffusion pathways in styloconic sensilla on the maxillary galea of larval Mamestra configurata (Walker) (Lepidoptera: Noctuidae) and five other species

The external ultrastructure of sensilla on the maxillary galea are investigated in Mamestra configurata and five other lepidopterous larvae using scanning electron microscopy. The galea and lacinia, comprising the mesal lobe of the maxilla, are either completely separate, fused, or incompletely fused in these species. The distal surface of the mesal lobe of all species examined bears two styloconic sensilla, three basiconic sensilla, and three trichoid sensilla, whereas the midventral wall of this lobe bears a campaniform sensillum. The latter sensillum is visible in only three of the six species examined. The styloconic and basiconic sensilla occupy a ventro anterior location, whereas the trichoid sensilla are positioned dorsoposteriorly. Interspecific comparisons of galeal size, as well as sensillar size, shape, and position are made for all species. The styloconic sensilla are the only sensillar type permeable to an aqueous solution of cobalt chloride when viewed by brightfield light microscopy in all species examined. Cobalt ions permeate through the terminal pore of each styloconic peg and percolate through the fenestrated fibrillar pore matrix, located directly below the pore. These ions permeate along the dendritic channel and accumulate in the adjacent sensillar sinus surrounding the peg and/or style by way of a presumably permeable dendritic sheath in all species, but to varying extents. The cuticular sidewall pores surrounding the terminal pore also appear to be permeable to cobalt ions in all the species examined. In most species examined, the styloconic sensilla are only minimally permeable to mercury ions. In these species, mercury ions permeate through the terminal pore, but become trapped within the plug of fenestrated fibrils within it. The sidewall pores are not permeable to mercury ions in any of the species examined. The styloconic sensilla are not permeable to lead ions in M. configurata or Malacosoma lutescens, the only species tested. © 1996 Wiley-Liss, Inc.     

Wg signaling via Zw3 and mad restricts self-renewal of sensory organ precursor cells in Drosophila

It is well known that the Dpp signal transducer Mad is activated by phosphorylation at its carboxy-terminus. The role of phosphorylation on other regions of Mad is not as well understood. Here we report that the phosphorylation of Mad in the linker region by the Wg antagonist Zw3 (homolog of vertebrate Gsk3-β) regulates the development of sensory organs in the anterior-dorsal quadrant of the wing. Proneural expression of Mad-RNA interference (RNAi) or a Mad transgene with its Zw3/Gsk3-β phosphorylation sites mutated (MGM) generated wings with ectopic sensilla and chemosensory bristle duplications. Studies with pMad-Gsk (an antibody specific to Zw3/Gsk3-β-phosphorylated Mad) in larval wing disks revealed that this phosphorylation event is Wg dependent (via an unconventional mechanism), is restricted to anterior-dorsal sensory organ precursors (SOP) expressing Senseless (Sens), and is always co-expressed with the mitotic marker phospho-histone3. Quantitative analysis in both Mad-RNAi and MGM larval wing disks revealed a significant increase in the number of Sens SOP. We conclude that the phosphorylation of Mad by Zw3 functions to prevent the self-renewal of Sens SOP, perhaps facilitating their differentiation via asymmetric division. The conservation of Zw3/Gsk3-β phosphorylation sites in vertebrate homologs of Mad (Smads) suggests that this pathway, the first transforming growth factor β-independent role for any Smad protein, may be widely utilized for regulating mitosis during development.     

Fine structure and distribution of antennal sensilla of the desert locust, Schistocerca gregaria (Orthoptera: Acrididae)

The fine structure and distribution of various types of antennal sensilla in three nymphal stages and in adults of both solitary-reared (solitary) and crowd-reared (gregarious) phases of the desert locust, Schistocerca gregaria, were investigated by scanning and transmission electron microscopy. Four types of sensilla were identified: sensilla basiconica, s. trichodea, s. coeloconica and s. chaetica. S. basiconica contain up to 50 sensory neurons, each of which displays massive dendritic branching. The sensillar wall is penetrated by a large number of pores. In contrast, s. trichodea contain one to three sensory neurons that branch to give five or six dendrites in the sensillar lumen; the sensillum wall is penetrated by relatively few pores. The s. coeloconica are situated in spherical cuticular pits on the antennal surface. The s. coeloconica are of two types: one type contains one to three sensory neurons with double sensillar walls penetrated by slit-like pores, whereas the second type contains four sensory neurons with non-porous double sensillar walls. The s. chaetica have a flexible socket and a thick non-porous sensillum wall and contain four sensory neurons that send unbranched dendrites to a terminal pore. A fifth sensory neuron of the s. chaetica terminates in a tubular body at the base of the hair. S. basiconica and coeloconica are normally distributed over the entire antennal flagellum, with a concentration in the middle segments; s. trichodea have three areas of concentration on the 5th, 10th and 14th flagellar segments. Sensilla chaetica are most abundant on the terminal segment. Locusts raised in solitary conditions have more olfactory sensilla (s. basiconica and s. coeloconica) than crowd-reared locusts. The difference in sensillar numbers is more evident in adults than in nymphs. These results suggest that differences in the odour-mediated behaviour of nymphs and adults, and between the phases of S. gregaria, may be attributable to differences at the sensory input level.