R. Naresh Singh

Sensilla on the tarsal segments and mouthparts of adult Drosophila melanogaster Meigen (Diptera, Drosophilidae)

Abstract Sensilla on the tarsal segments of legs and on the external and internal mouthparts of adult Drosophila melanogaster Meigen (Diptera : Drosophilidae) were studied by light and electron microscopy. Tarsi are invested with bracteate and bractless bristles. Ethanolic silver nitrate stain distinguishes dendrite-containing bristles from other hairs on the tarsi. Stained bristles are bractless. Males have more silver-staining bristles than females on the I st 4 tarsal segments of prothoracic legs. There are 33 – 34 taste bristles on each half of the labellum. Many of them are 2-pronged. Both prongs have a pore at the tip, but chemosensory dendrites enter only in one of them. The taste bristles in the 2 medial rows on each half of the labellum are predominantly associated with 4 chemosensory neurons and the peripheral bristles usually contain 2 chemosensory neurons each. Five groups of paired sense organs are known to exist on the internal mouthparts. Amongst 9 sensilla present in the labral sense organ, electron microscopy shows that sensilla numbers 1 – 6 have one mechanosensory neuron each. Sensillum number 7 has 8 neurons with dendrites arranged in 3 groups: 2 triplets and a pair. Sensilla 8 and 9 have 2 neurons each. The ventral cibarial sense organ (VCSO) has 2 sensilla, one with 2 and the other with 4 neurons. Rows of hollow satellite bristles forming 2 sensory systems, dorsal and ventral to the VCSO have one mechanosensory neuron each. The dorsal cibarial sense organ has 2 sensilla on either side of the midline, each containing 3 neurons. The paper describes the fine structural morphology of taste receptors and enumerates axons of all the prominent nerves of mouthparts at different levels. An external nerve loop connecting directly 2 regions of the brain, the tritocerebrum and suboesophageal ganglion, is reported.

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

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

Fine structure of the sensory organs of Drosophila melanogaster Meigen larva (Diptera: Drosophilidae)

Abstract Fine structure of the prominent external and internal sensory organs of Drosophila melaogaster Meigan (Diptera : Drosophilidae) larva was determined by transmission electron microscopy (TEM). The external sensory organs, namely, antennal, maxillary, ventral and labial organs, dorsal pits, sensory cones on the 8th and 9th abdominal segments, and the sensory hairs on the body of the larva, were studied. A new knob in pit (KIP) sensillum innervated by 3 dendrites was found on the dorsolateral surface on either side. Four tufts of hairs at the posterior end of each great lateral tracheal trunk were found to be sensory with a dendrite at the base of each tuft. Internally in the pharynx of the larva, 3 groups of symmetrically located sensory organs, namely, the anteroventral, dorsal, and posteroventral groups were found. In all they contain 10 pairs of sensilla. Amongst them, 2 pairs of sensilla were found to be of the compound type; one having 9 dendrites arranged in 3 groups of 4, 3 and 2; while the other had 6 dendrites grouped as 2 and 4. In addition, 2 groups of sensilla were found on the internal dorsal fold on either side. Similarities were observed in the dendritic organisation of sensilla in the internal mouthparts of the Drosophila larva and the adult. Unlike nerves of the adult, the larval nerves connected with the dorsal and maxillary organs have a significantly thick layer of glial cells ensheathing the bundle of axons.

Neurobiology of the gustatory systems of Drosophila and some terrestrial insects

Insects have been favorites for the study of taste perception in the last few decades. They have been used for anatomical, behavioral, developmental, genetic, and physiological studies related to gustation and feeding response. Several genes known to affect the formation of gustatory sensilla or alter the feeding behavior of insects such as Drosophila are known. Studies related to signal transduction, coding of gustatory information, and the nature and constitution of genes involved in taste perception have also been taken up with insects in recent years. The understanding of basic mechanisms of taste perception in insects is likely to lead to better management of useful as well as harmful insects. Microsc. Res. Tech. 39:547–563, 1997.

Sensilla on the third antennal segment of Drosophila melanogaster meigen (Diptera : Drosophilidae)

Abstract The distribution and morphology of the sensilla on the 3rd antennal segment of Drosophila melanogaster Meigen (Diptera : Drosophilidae) were studied with light and electron microscopy. Four types of hairs were identified. Three types of hairs innervated by dendrites are sensilla basiconica, sensilla coeloconica and sensilla trichodea. They occur amongst a large number of the 4th type of uninnervated hairs or spinules. Sensilla basiconica and coeloconica can be easily identified by light microscopy on staining with 0.1016 silver nitrate in 70% ethanol. The tips of sensilla basiconica and coeloconica appear dark brown. Most of the sensilla trichodea and spinules remain unstained. Sensilla basiconica conform to the single-walled, multiporous sensilla, having poretubules and branched dendrites. Sensilla coeloconica are double-walled and have longitudinal channels near the tip. No wall pores are found on sensilla trichodea. Dendrites do not branch in sensilla coelonica and trichodea. A mechanosensory dendrite with characteristic tubular body is absent in these sensilla. Populations of sensilla basiconica and sensilla trichodea occur in diametrically opposite, distinct regions on the 3rd antennal segment-the former in the dorsomedial and the latter in the ventrolateral regions, whereas sensilla coeloconica are distributed on most of the anterior and posterior surfaces, including the cavity walls of the sacculus. The axons are arranged in distinct groups in the antennal nerves at the stalk of the 3rd segment. This grouping becomes more pronounced in the nerve prior to its entry into the brain.

Fine structure and primary sensory projections of sensilla located in the sacculus of the antenna of Drosophila melanogaster

Sensilla lining the inner walls of the sacculus on the third antennal segment of Drosophila melanogaster were studied by light and transmission electron microscopy. The sacculus consists of three chambers: I, II and III. Inside each chamber morphologically distinct groups of sensilla having inflexible sockets were observed. Chamber I contains no-pore sensilla basiconica (np-SB). The lumen of all np-SB are innervated by two neurons, both resembling hygroreceptors. However, a few np-SB contain one additional neuron, presumed to be thermoreceptive. Chamber II houses no-pore sensilla coeloconica (np-SC). All np-SC are innervated by three neurons. The outer dendritic segments of two of these neurons fit tightly to the wall of the lumen and resemble hygroreceptor neurons. A third, more electron-dense sensory neuron, terminates at the base of the sensillum and resembles a thermoreceptor cell. Chamber III of the sacculus is divided into ventral and dorsal compartments, each housing morphologically distinct grooved sensilla (GS). The ventral compartment contains thick GS1, and the dorsal compartment has slender sensilla GS2. Ultrastructurally, both GS1 and GS2 are doublewalled sensilla with a longitudinal slit-channel system and are innervated by two neurons. The dendritic outer segment of one ofthe two neurons innervates the lumen of the GS and branches. On morphological criteria, we infer this neuron to be olfactory. The other sensory neuron is probably thermoreceptive. Thus, the sacculus in Drosophila has sensilla that are predominantly involved in hygroreception, thermoreception, and olfaction. We have traced the sensory projections of the neurons innervating the sacculus sensilla of chamber III using cobaltous lysine or ethanolic cobalt (II) chloride. The fibres project to the antennal lobes, and at least four glomeruli (VM3, DA3 and DL2-3) are projection areas of sensory neurons from these sensilla. glomerulus DL2 is a common target for the afferent fibres of the surface sensilla coeloconica and GS, whereas the VM3, DA3 and DL3 glomeruli receive sensory fibres only from the GS.

Primary sensory projections from the labella to the brain of Drosophila melanogaster Meigen (Diptera : Drosophilidae)

Abstract Golgi silver impregnation of sensory neurons arising from labellar taste sensilla of Drosophila melanogaster Meigen (Diptera : Drosophilidae) revealed 7 distinct types I-VII of primary (sensory) fibres projecting to the suboesophageal ganglion (SOG) of the brain. Each fibre was classified on the bases of the neuropil volume occupied by its terminal arborisation, the shape of neuropil region receiving the arborisations and the detailed morphology of the arborisations. The primary sensory fibre projections from the labella are confined to the SOG where they project mainly in the anterior and central neuropils. No labellar sensory fibres project to posterior SOG. Of these 7 types of sensory fibres, three (III, IV and VII) show ipsilateral projections, while others have both ipsi-, and contralateral branches. Four types of interneurons are suggested to be associated with taste perception. Type A interneurons are local interneurons with arborisations confined only to the taste sensory neuropil of the SOG. The types B - D interneurons are interganglionic/output neurons with axons projecting to various brain regions-SOG, calyces of the mushroom bodies, tritocerebrum and thoracic ganglia. These projections suggest that more than one centre (SOG, tritocerebrum, calyces of the mushroom bodies and thoracic ganglia) are involved in processing gustatory information.

Functional implications of the projections of neurons from individual labellar sensillum of Drosophila melanogaster as revealed by neuronal-marker horseradish peroxidase

SummaryEarlier studies using Golgi silver impregnations from the labellar sensilla of adult Drosophila melanogaster revealed seven types of sensory axons projecting into the suboesophageal ganglion of the brain. These sensory terminals were designated as coiled fibres (type-I), shrubby fibres (type-II), ipsilateral ventral fibres (type-III), ipsilateral dorsal fibres (type-IV), contralateral ventral fibres (type-V), contralateral dorsal fibres (type-VI), and central fibres (type-VII). The present study identifies the projections of sensory neurons present in a single labellar taste-sensillum, using the neuronal marker horseradish peroxidase (HRP). Although the taste sensillum in question has five neurons, in a given experiment only one or at the most two neurons are labelled. The type of neuron labelled was usually specific to the stimulant solute (sucrose, sodium chloride or potassium chloride) present in the HRP solution. Although type-II fibres get labelled most of the time, irrespective of the stimulant present in HRP solution, type-IV fibres are labelled when attractants (0.1 M sucrose or ≤0.1 M sodium chloride) are used as stimulants in HRP solution. Type-VI fibres are labelled when the stimulant is 0.1 M potassium chloride, a repellent. HRP dissolved in distilled water revealed type-I coiled fibres. Besides revealing projections of sensillar neurons to the brain the present technique also inferred their possible function. Incubation of whole-brain tissue with 0.04% 3,3′-diaminobenzidine tetrahydrochloride in presence of 0.06% hydrogen peroxide suggested that the glomerular organization is also present in the taste-sensory region as it is in olfactory neuropile.

Ultrastructure of the femoral chordotonal organs and their novel synaptic organization in the legs of Drosophila melanogaster Meigen (Diptera : Drosophilidae)

Abstract The chordotonal or scolopophorous organs located in the femoral segment of all the legs of both male and female Drosophila melanogaster (Diptera : Drosophilidae) were examined by light and transmission electron microscopes. The femoral chordotonal organs (FCO) are arranged in 3 groups: one large group consists of about 32 aligned scolopes whose distal ends extend and terminate at the distal epicuticular surface of the femur; the other 2 central groups together contain about 42 scattered scolopes, and distally they terminate into the membranes connecting the muscles of the femur. No sexual dimorphism is evident either in the numbers of scolopidial groups or the total number of scolopidia in both sexes. Each scolopidium is innervated by 2 neurons of which one is less electron-dense than the other. The dendrites of these neurons bear sensory cilia. The fine structure of these chordotonal sensilla suggests that they probably respond to stretch or flexion. Proximally in the femur, the axons from FCO form a novel glomerular organization. These axons show lateral extensions and form different morphological types of synapses among themselves. From the presence of a large number of putative chemical synapses in the legs, it is evident that some degree of information processing is taking place in D. melanogaster at the periphery before being relayed to the central nervous system.

Sensory organs on the body parts of the bed-bug Cimex hemipterus fabricius (Hemiptera : Cimicidae) and the anatomy of its central nervous system

Abstract Anatomy of the sensory organs on the prominent body parts of the adult bed-bug Cimex hemipterus (Hemiptera: Cimicidae) and its central nervous system (CNS) was studied by light, transmission, or scanning electron microscopy. The distal tips of antenna and rostrum were found to have rich complements of sensilla. The antenna has both olfactory and gustatory sensilla. Olfactory sensilla project to the antennal lobe organized in the form of glomeruli, while the 2nd component, presumably from gustatory sensilla, projects to the suboesophageal ganglion. The ultrastructure of the sensory pegs on the rostrum of C. hemipterus does not resemble the chemosensilla of adult insects; rather they resemble the larval sensilla of Drosophila melanogaster in the maxillary organ. Earlier we believed this to be a gustatory organ. A few similar sensilla also occur on the antenna, indicating its multimodal role. Amongst the 3 types of sensory hairs located on legs, there are only a few gustatory hairs (7–10 hairs) on the tibia. The pointed and serrate mechanosensory hair types occur in abundance; the serrate type are prominently present on the lateral surface of the legs. On other parts of the body such as the thorax or abdomen, serrate hairs are most abundant. Both the distal segment of antenna and rostrum are invested by 2 nerves, where the axon counts of the 2 antennal nerves are 380 and 425, while each rostral nerve on average has 205 axons. Abundant clusters of microtubules were found in the brain, thoracio-abdominal ganglia, leg-nerves, and the space between muscles and cuticle. These conspicuous microtubule-clusters occur in interaxonal space, mainly glial cells, in the nervous system. In addition, the glial cells have osmiophilic junctions amongst themselves. A novel “hinge and joint” system, which controls the cross-section of the food canal and the salivary duct in an inversely related manner, was found in the rostrum of the bed-bug.