Dual, multilayered somatosensory maps formed by antennal tactile and contact chemosensory afferents in an insect brain

The antennae of most insects move actively and detect the physical and chemical composition of objects encountered by using their associated tactile sensors. Positional information is required for these sensory modalities to interpret the physical environment. Although we have a good understanding of antennal olfactory pathways, little is known about the destinations of antennal mechanosensory and contact chemosensory (gustatory) receptor neurons in the central nervous system. The cockroach Periplaneta is equipped with a pair of long, thin antennae, which are covered in bristles. The distal portions of each antenna possess about 6,500 bimodal bristles that house one tactile sensory and one to four contact chemosensory neurons. In this study, we investigated the morphologies of bimodal bristle receptor afferents by staining individual or populations of bristles. Unlike olfactory afferents, which project exclusively into the glomeruli in the ventral region of the deutocerebrum, both the presumptive mechanosensory and the contact chemosensory afferents projected into the posterior dorsal region of the deutocerebrum and the anterior region of the subesophageal ganglion. Each afferent showed multilayered segmentation and spatial occupation reflecting its three-dimensional position in the periphery. Presumptive contact chemosensory afferents, characterized by their thin axons and unique branching pattern, occupied more medioventral positions compared with the presumptive tactile afferents. Furthermore, projection fields of presumptive contact chemosensory afferents from single sensilla tended to be segregated from each other. These observations suggest that touch and taste positional information from the antenna is precisely represented in primary centers in a modality-specific manner.     

Modality-specific axonal projections in the CNS of the flies Phormia and Drosophila

There is a rich history of behavioral and physiological studies on the leg sensory systems of flies. Here we examine the anatomy of the sensory axons of two species of fly and demonstrate that the location of the axonal projections in the CNS can be correlated with the modality they encode. We studied receptors associated with proprioceptive, tactile, and multimodal hairs. Proprioceptive hairs occur in clusters, called hair plates, and are situated near joints. The neuron innervating each proprioceptive hair has a large axon and coarse arborization in the intermediate neuropil. Tactile receptors have smaller arbors, which are located in a ventral region of the thoracic neuromere. Finally, the multimodal hairs are each innervated by one tactile and four chemosensory neurons. The single tactile neuron has a central arbor that is indistinguishable from those of the tactile hairs; the four chemosensory neurons project to yet a third region of neuropil near the ventral surface of each neuromere. Thus there is a clear modality-specific segregation of axonal arbors in the CNS. This organization is identical in Phormia and Drosophila and thus apparently highly conserved within the Diptera. We presume that, as in other insect sensory systems, this anatomical specificity is linked to synaptic specificity.     

Neural control of leg movements in a metamorphic insect: sensory and motor elements of the larval thoracic legs in Manduca sexta

During the metamorphosis of the hawkmoth Manduca sexta the larval thoracic legs degenerate to be replaced in the adult by legs of very different form and function. This change must be accompanied by a reorganization of the neural circuits controlling leg movements. As an initial step in the study of this reorganization we describe here the sensory and motor elements of this circuitry in the larval stage of life. Sensory neurons innervating mechanoreceptive hairs on the thoracic surface were stained individually with cobalt. Those innervating hairs on the general thoracic surface project topographically into two ventral regions of the segmental ganglia. Sensory neurons innervating leg sensilla also map topographically to the more ventral of these regions but in addition have arborizations in a midlateral region. The density of branching within this lateral "leg neuropil" is greatest for sensory neurons form sensilla on the more distal leg segments. Leg motor neurons were identified with intracellular recording and cobalt injection techniques. Those innervating muscles controlling distal leg segments have dense dendritic arbors in the lateral "leg neuropil," while motor neurons controlling more proximal segments and muscles of the ventral body wall have extensive arborizations in a dorsomedial region of the ganglion. In general, flexor motor neurons are excited by medial and inhibited by lateral leg sensilla, while the opposite is true of extensors. Distal segment motor neurons respond most strongly to sensory neurons from distal segments, thus suggesting some interaction within the lateral "leg neuropil." Thus, in the larval nervous system a highly ordered array of of sensory and motor elements underlies the specific behavioral responses of the legs to tactile stimulation.     

Correlation between the receptive fields of locust interneurons,their dendritic morphology,and the central projections of mechanosensory neurons

The relationships between the morphology and receptive fields of local and intersegmental interneurons that process mechanosensory information from a hindleg of the locust have been analysed. Sensory neurons from tactile hairs project to ventral areas of neuropil in the metathoracic ganglion where they form a 3-dimensional somatotopic map of a hindleg. By contrast, sensory neurons from a proprioceptor at the femoro-tibial joint (the femoral chordotonal organ) project to lateral and more intermediate areas of neuropil and have no branches in the most ventral regions of neuropil. Particular local and intersegmental interneurons respond to stimulation of specific arrays of hairs on a hindleg, or to movements of particular joints. Their receptive fields are defined, in part, by the patterns of excitatory, monosynaptic connections made by these afferents. Each interneuron has a characteristic receptive field and a characteristic morphology defined by its array of branches in the regions of neuropil containing the projections of the afferents that provide its monosynaptic inputs. Interneurons with inputs exclusively from tactile hairs have branches in the most ventral regions of neuropil, while those with exclusively proprioceptive inputs have branches only in more intermediate levels of neuropil. Interneurons with extensive receptive fields from tactile hairs also have extensive areas of branching within the ventral neuropil. Interneurons with receptive fields restricted to particular regions of the leg have branches restricted to the ventral region of neuropil containing the projections of afferents from that part of the leg. Thus, interneurons with inputs only from hairs on the tarsus have branches in the posterior region of neuropil corresponding to the projections of the tarsal afferents, while interneurons with receptive fields on the femur have branches in more anterior regions of neuropil corresponding to the projections of the femoral hair afferents. Interneurons with receptive fields on the tibia have branches in neuropil between the tarsal and femoral projections. © 1993 Wiley-Liss, Inc.     

Sensillum-specific, topographic projection patterns of olfactory receptor neurons in the antennal lobe of the cockroach Periplaneta americana

In vertebrates and many invertebrates, olfactory signals detected by peripheral olfactory receptor neurons (ORNs) are conveyed to a primary olfactory center with glomerular organization in which odor-specific activity patterns are generated. In the cockroach, Periplaneta americana, ORNs in antennal olfactory sensilla project to 205 unambiguously identifiable antennal lobe (AL) glomeruli that are classified into 10 glomerular clusters (T1-T10 glomeruli) innervated by distinct sensory tracts. In this study we employed single sensillum staining techniques and investigated the topographic projection patterns of individual ORNs to elucidate the relationship between sensillum types and glomerular organization in the AL. Axons of almost all ORNs projected to individual glomeruli. Axons of ORNs in perforated basiconic sensilla selectively innervated the anterodorsal T1-T4 glomeruli, whereas those in trichoid and grooved basiconic sensilla innervated the posteroventral T5-T9 glomeruli. About 90% of stained ORNs in trichoid sensilla sent axons to the T5 glomeruli and more than 90% of ORNs in grooved basiconic sensilla innervated the T6, T8, and T9 glomeruli. The T5 and T9 glomeruli exclusively receive sensory inputs from the trichoid and grooved basiconic sensilla, respectively. All investigated glomeruli received convergent input from a single type of sensillum except F11 glomerulus in the T6 glomeruli, which was innervated from both trichoid and grooved basiconic sensilla. These results suggest that ORNs in distinct sensillum types project to glomeruli in distinct glomerular clusters. Since ORNs in distinct sensillum types are each tuned to distinct subsets of odorant molecules, the AL is functionally compartmentalized into groups of glomeruli.     

Hypophysiotropic neurons of the paraventricular nucleus respond in spatially, temporally, and phenotypically differentiated manners to acute vs. repeated restraint stress: rapid publication

The antennules of decapod crustaceans are covered with thousands of chemosensilla that mediate odor discrimination and orientation behaviors. Most studies on chemoreception in decapods have focused on the prominent aesthetasc sensilla. However, previous behavioral studies on lobsters following selective sensillar ablation have revealed that input from nonaesthetasc antennular chemosensilla is sufficient for many odor-mediated behaviors. Our earlier examination of the setal types on the antennules of the Caribbean spiny lobster Panulirus argus revealed three types of nonaesthetasc chemosensilla. The most abundant and widely distributed of these is the hooded sensillum. The present study describes the detailed ultrastructure of antennular hooded sensilla and the physiological response properties of their receptor neurons. Light and scanning and transmission electron microscopy were used to examine structural characteristics, and electrophysiology was used to examine single-unit responses elicited by focal chemical and mechanical stimulation of antennular hooded sensilla. Hooded sensilla have a porous cuticle and are innervated by 9-10 chemosensory and 3 mechanosensory neurons whose dendrites project to the distal end of the sensillum. Hooded sensillar chemosensory neurons responded to waterborne chemicals, were responsive to only one of the six tested single compounds, and had different specificities. Hooded sensillar mechanosensory neurons were not spontaneously active. They had low sensitivity in that they responded to tactile but not waterborne vibrations, and they responded to sensillar deflection with phasic bursts of activity. These results support the idea that hooded sensilla are bimodal chemo-mechanosensilla and are receptors in an antennular chemosensory pathway that parallels the well-described aesthetasc chemosensory pathway.     

Morphology and somatotopic organisation of the central projections of afferents from tactile hairs on the hind leg of the locust.

The morphology and organisation of the central projections of tactile hair afferents from the hind leg of the locust, Schistocerca gregaria, were examined by staining individual hair afferents. Each tactile hair on the femur, tibia, and tarsus is innervated by a single sensory afferent, which projects to the ipsilateral half of the metathoracic ganglion. Afferents arborize in the ventralmost and lateral ventral association centres (vVAC and lVAC). The projections are organised somatotopically in a map with three axes, according to the position of the hair on the leg. First, proximo-distal: afferents from hairs on the proximal leg segments project more anteriorly than do those from hairs on distal leg segments. Moreover, on any given segment the afferents from the more proximal hairs project more anteriorly than do the afferents from the distal hairs. Second, antero-posterior: afferents from hairs on the posterior surface of the leg project more medially than do afferents from anterior hairs. Third, dorso-ventral: afferents from hairs on ventral parts of the leg project more ventrally than do afferents from the dorsal hairs. The afferents from posterior and anterior hairs project to an area between the central projections from dorsal hairs and ventral hairs. The position of a projection within the map is dependent upon the location of the hair on the leg and not the peripheral routes taken by the axon of its afferent to reach the ganglion.     

Sensory neuron development revealed by taurine immunocytochemistry in the honeybee

The formation of ommatidia in the compound eyes and sensilla on the antennae of the honeybee was followed and the development of their sensory neurons was traced using an antiserum against taurine as a marker. Taurine-like immunoreactivity (Tau-IR) is expressed in sensory neurons of several modalities, namely visual, olfactory, gustatory, and mechanosensory. Staining intensity is very high in the larva and in the first half of the pupal stage and gradually decreases towards the end of metamorphosis. In the photoreceptor cells of the compound eyes, Tau-IR can be detected from the fifth larval instar onwards, prior to differentiation of other components of the ommatidium. Already in the midstage larvae, when the antennal primordia of the adult still lie within the peripodial cavity, a few presumably mechanosensory neurons are labelled in the pedicellus of the developing antenna. The majority of the antennal sensory neurons which are located on the flagellum start to exhibit Tau-IR upon pupation, long before any cuticular specializations such as sensory hairs or plates are detectable. All known types of antennal sensilla were identified and it could be shown that all of them are innervated by Tau-IR sensory neurons. Thus, taurine immunocytochemistry can be applied as a useful label for developing sensory neurons. Functional implications of taurine during development are discussed. © 1995 Wiley-Liss, Inc.     

Serotonin‐immunoreactive sensory neurons in the antenna of the cockroach Periplaneta americana

The antennae of insects contain a vast array of sensory neurons that process olfactory, gustatory, mechanosensory, hygrosensory, and thermosensory information. Except those with multimodal functions, most sensory neurons use acetylcholine as a neurotransmitter. Using immunohistochemistry combined with retrograde staining of antennal sensory neurons in the cockroach Periplaneta americana, we found serotonin‐immunoreactive sensory neurons in the antenna. These were selectively distributed in chaetic and scolopidial sensilla and in the scape, the pedicel, and first 15 segments of the flagellum. In a chaetic sensillum, A single serotonin‐immunoreactive sensory neuron cohabited with up to four serotonin‐negative sensory neurons. Based on their morphological features, serotonin‐immunopositive and ‐negative sensory neurons might process mechanosensory and contact chemosensory modalities, respectively. Scolopidial sensilla constitute the chordotonal and Johnston's organs within the pedicel and process antennal vibrations. Immunoelectron microscopy clearly revealed that serotonin‐immunoreactivities selectively localize to a specific type of mechanosensory neuron, called type 1 sensory neuron. In a chordotonal scolopidial sensillum, a serotonin‐immunoreactive type 1 neuron always paired with a serotonin‐negative type 1 neuron. Conversely, serotonin‐immunopositive and ‐negative type 1 neurons were randomly distributed in Johnston's organ. In the deutocerebrum, serotonin‐immunoreactive sensory neuron axons formed three different sensory tracts and those from distinct types of sensilla terminated in distinct brain regions. Our findings indicate that a biogenic amine, serotonin, may act as a neurotransmitter in peripheral mechanosensory neurons. J. Comp. Neurol. 522:414–434, 2014. © 2013 Wiley Periodicals, Inc.     

Somatotopic mapping of sensory neurons innervating mechanosensory hairs on the larval prolegs of Manduca sexta

The abdominal prolegs are the principal locomotory appendages of the larval tobacco hornworm, Manduca sexta. The prolegs bear numerous mechanosensory bristle sensilla, each innervated by an afferent neuron that arborizes within the central nervous system (CNS). Based on their positions on the proleg, we have divided the sensilla into planta hairs (PHs), lateral hairs (LHs), and medial hairs (MHs). Previously, we found that PH afferents produce monosynaptic excitatory postsynaptic potentials (EPSPs) in proleg retractor muscle motoneurons, the size of which depends on the position of the hair in the PH array. In this paper we examined the central arbors of the proleg afferents to determine whether there was an anatomical correlate to the pattern of synaptic strengths. We found that the afferent arbors are arranged somatotopically within the CNS in a pattern similar to that for bristle afferents elsewhere on the abdomen; i.e., the anterior-posterior and medial-lateral position of a hair on the proleg was reflected in the location of the afferent arbor along the corresponding axes within sensory neuropil. All afferents terminated within a similar ventral region of neuropil. The arbors of PH, MH, and to a lesser extent, LH afferents, were enlarged as compared to afferents innervating hairs elsewhere on the abdomen. This feature, combined with the dense innervation of the proleg, causes the proleg region to be relatively overrepresented in sensory neuropil. We also examined the afferents innervating a pair of ventral midline hairs (VMHs) present in each abdominal segment, which, unlike the other afferents, showed segment-specific central arbors. We conclude that the somatotopic mapping of afferent arbors may contribute to the specificity of synaptic connections in this system.     

Sex‐specific antennal sensory system in the ant Camponotus japonicus: Glomerular organizations of antennal lobes

Ants have well‐developed chemosensory systems for social lives. The goal of our study is to understand the functional organization of the ant chemosensory system based on caste‐ and sex‐specific differences. Here we describe the common and sex‐specific glomerular organizations in the primary olfactory center, the antennal lobe of the carpenter ant Camponotus japonicus. Differential labeling of the two antennal nerves revealed distinct glomerular clusters innervated by seven sensory tracts (T1–T7 from ventral to dorsal) in the antennal lobe. T7 innervated 10 glomeruli, nine of which received thick axon terminals almost exclusively from the ventral antennal nerve. Coelocapitular (hygro‐/thermoreceptive), coeloconic (thermoreceptive), and ampullaceal (CO₂‐receptive) sensilla, closely appositioned in the flagellum, housed one or three large sensory neurons supplying thick axons exclusively to the ventral antennal nerve. These axons, therefore, were thought to project into T7 glomeruli in all three castes. Workers and virgin females had about 140 T6 glomeruli, whereas males completely lacked these glomeruli. Female‐specific basiconic sensilla (cuticular hydrocarbon‐receptive) contained over 130 sensory neurons and were completely lacking in males' antennae. These sensory neurons may project into T6 glomeruli in the antennal lobe of workers and virgin females. Serotonin‐immunopositive neurons innervated T1–T5 and T7 glomeruli but not T6 glomeruli in workers and virgin females. Because males had no equivalents to T6 glomeruli, serotonin‐immunopositive neurons appeared to innervate all glomeruli in the male's antennal lobe. T6 glomeruli in workers and virgin females are therefore female‐specific and may have functions related to female‐specific tasks in the colony rather than sexual behaviors. J. Comp. Neurol. 518:2186–2201, 2010. © 2010 Wiley‐Liss, Inc.     

Two closely located areas in the suboesophageal ganglion and the tritocerebrum receive projections of gustatory receptor neurons located on the antennae and the proboscis in the moth Heliothis virescens

Sucrose stimulation of gustatory receptor neurons on the antennae, the tarsi, and the mouthparts elicits the proboscis extension reflex in many insect species, including lepidopterans. The sensory pathways involved in this reflex have only partly been investigated, and in hymenopterans only. The present paper concerns the pathways of the gustatory receptor neurons on the antennae and on the proboscis involved in the proboscis extension reflex in the moth Heliothis virescens (Lepidoptera; Noctuidae). Fluorescent dyes were applied to the contact chemosensilla, sensilla chaetica on the antennae, and sensilla styloconica on the proboscis, permitting tracing of the axons of the gustatory receptor neurons in the central nervous system. The stained axons showed projections from the two appendages in two closely located but distinct areas in the suboesophageal ganglion (SOG)/tritocerebrum. The projections of the antennal gustatory receptor neurons were located posterior-laterally to those from the proboscis. Electrophysiological recordings from the receptor neurons in s. chaetica during mechanical and chemical stimulation were performed, showing responses of one mechanosensory and of several gustatory receptor neurons. Separate neurons showed excitatory responses to sucrose and sinigrin. The effect of these two tastants on the proboscis extension reflex was tested by repeated stimulations with solutions of the two compounds. Whereas sucrose elicited extension in 100% of the individuals in all repetitions, sinigrin elicited extension in fewer individuals, a number that decreased with repeated stimulation.     

Mechanosensory pegs constitute stridulatory files in grasshoppers.

Stridulatory files on the inner face of hindleg femora were shown to consist of mechanosensory pegs in males and females of Syrbula montezuma (Saussure) and in males of Chorthippus biguttulus (L.). Females of Chorthippus had stiff protuberances on their stridulatory files, with an innervated tubercle instead of pegs. Pegs and tubercles of adult grasshoppers were shown to develop from innervated tubercular hairs present from the first instar onward in Chorthippus. In adults of Chorthippus, two sensory cells innervated each peg of males and each tubercle of females. Central projections of these afferents from the stridulatory files were very similar to those of the neighboring tactile hairs on the femur. The afferents from pegs in Syrbula responded to deflection and pressure introduced via the widened cuticular cap. In both species, selective stimulation of femoral cuticular receptors elicited antagonistic reflex responses in a coxal retractor muscle: pegs inhibited and neighboring hairs raised the efferent tonic discharges. Apparently, in these two distantly related grasshopper species, stridulatory files function as both sound-producing and proprioceptive organs.     

Central projections of axons from taste hairs on the labellum and tarsi of the blowfly, Phormia regina Meigen.

Taste hairs are located on the labellum and tarsi of blowflies. These multimodal hairs consist of four functionally distinct chemoreceptors and a mechanoreceptor. By staining selected multimodal hairs, we sought to identify the central projection patterns of multiple and single axons from those hairs. On each side of the labellum there are 11 "largest" hairs (LH). The neurons associated with the anteriormost (LH-1), posteriormost (LH-11), and one lateral (LH-6) hair on the labellum were stained selectively with cobaltous sulfide. The overall projection pattern in the central nervous system (CNS) for axons from LH-1 and LH-11 is similar and differs markedly from axons from LH-6. At least three individual axon-projection patterns were determined for each labellar hair filled, indicating a partial functional organization for axons from multimodal hairs. One identified axon, the dorsalmost axon, has terminal arborizations that do not differ with the location of its associated hair. Another axon, thicker than the others, projects to a region that is distinct from the four thin axons. Within this region the arborizations of the thick axons occupy different areas depending on the location of their associated hair. Neurons from the largest hairs on the distalmost tarsomere (D5) of each leg were also stained and consisted of one thick and four thin axons. All axons except one thin axon from tarsal D5 hairs terminate in their respective leg neuromeres. The remaining thin axon projects to the suboesophageal ganglion ipsilateral to the hair filled and terminates in the same region as a branch of the labellar dorsalmost axon. These data suggest that axonal arbors from multimodal hairs have a limited functional and somatotopic organization in the blowfly CNS.     

Morphological analysis of the primary center receiving spatial information transferred by the waggle dance of honeybees

The waggle dancers of honeybees encodes roughly the distance and direction to the food source as the duration of the waggle phase and the body angle during the waggle phase. It is believed that hive‐mates detect airborne vibrations produced during the waggle phase to acquire distance information and simultaneously detect the body axis during the waggle phase to acquire direction information. It has been further proposed that the orientation of the body axis on the vertical comb is detected by neck hairs (NHs) on the prosternal organ. The afferents of the NHs project into the prothoracic and mesothoracic ganglia and the dorsal subesophageal ganglion (dSEG). This study demonstrates somatotopic organization within the dSEG of the central projections of the mechanosensory neurons of the NHs. The terminals of the NH afferents in dSEG are in close apposition to those of Johnston's organ (JO) afferents. The sensory axons of both terminate in a region posterior to the crossing of the ventral intermediate tract (VIT) and the maxillary dorsal commissures I and III (MxDCI, III) in the subesophageal ganglion. These features of the terminal areas of the NH and JO afferents are common to the worker, drone, and queen castes of honeybees. Analysis of the spatial relationship between the NH neurons and the morphologically and physiologically characterized vibration‐sensitive interneurons DL‐Int‐1 and DL‐Int‐2 demonstrated that several branches of DL‐Int‐1 are in close proximity to the central projection of the mechanosensory neurons of the NHs in the dSEG. J. Comp. Neurol. 521:2570–2584, 2013. © 2013 Wiley Periodicals, Inc.     

Compartments and the topography of leg afferent projections in Drosophila

The legs of Drosophila are covered with mechanosensory bristles, innervated by sensory neurons that project to the CNS in a very orderly manner. We examined this afferent projection by staining the sensory neurons associated with identified bristles in wild-type, engrailed and scute flies. We observe that anterior neurons project to an anterior region of the ventral neuropil, while posterior neurons project to a more posterior region. We rule out that this difference depends on the compartment of origin of the receptors. Our results also argue against explanations based on other factors that might correlate to anterior/posterior position: peripheral organization of the leg nerve, competitive interactions, or differences in times of birth. We suggest that position itself is the primary determinant of this projection.     

Synaptic Connections of Different Strength Between Wind-sensitive Hairs and an Identified Projection Interneuron in the Locust

An identified intersegmental interneuron in Locusta and Schistocerca, with its cell body in the fourth abdominal ganglion and an axon which projects to the brain is excited by mechanosensory inputs from receptors on the head and neck. The organization of its receptive field, the types of sensory receptors which contribute to it and the patterns and strengths of the afferent connections were investigated by intracellular recording from the axon of the interneuron close to a spike-initiating site in the prothoracic ganglion. The receptive field of the interneuron consists of a small patch of hairs on the head ipsilateral to the axon, and from hairs on two regions of the prosternum (a cuticular structure on the ventral surface of the prothoracic segment), first an ipsilateral, lateral region and second a medial but contralateral region. Hairs on the pronotum (dorsal neck) also contribute but were not investigated here. Each spike in the afferent from a hair with a filiform appearance and with a pigmented base on the prosternum consistently evokes an EPSP in the interneuron. These have a short and constant latency, indicating that the connection is probably direct. The head hairs also appear to make direct connections with the interneuron in the prothoracic ganglion, so that the spike-initiating site here can integrate signals evoked by wind on the head and on the prosternum. Stiff tactile hairs on the prosternum do not connect with the interneuron. The EPSPs evoked by the long filiform hairs are consistently larger than those produced by the short filiform hairs and a single spike in some of the afferents from the long filiform hairs can evoke a spike in the interneuron. The effectiveness of an afferent is therefore correlated with the length of the filiform hair it innervates. The hairs with the most powerful effects are always the longest and occur in the same position on every locust. The shape of the receptive field and the different strengths of connections are apparent even in early larval instars. The axonal branches of the interneuron are restricted to the same side of the ganglion as the axon itself. Afferents from filiform hairs on the medial region of the prosternum project contralaterally, and those from the lateral region project ipsilaterally. Afferents from some of the head hairs project ipsilaterally directly to the prothoracic ganglion. The terminals of all these afferents overlap with the branches of the interneuron. By contrast, the afferents of tactile hairs which do not connect, project to different regions of neuropile. The connections ensure that the high sensitivity of the filiform hairs is maintained at the first stage in the central processing and suggest a role for this interneuron in supplying information about small changes in air currents that may be of use in controlling steering manoeuvres during flight.     

Birth of ophthalmic trigeminal neurons initiates early in the placodal ectoderm

In the primary olfactory center of animals, glomeruli are the relay stations where sensory neurons expressing cognate odorant receptors converge onto interneurons. In cockroaches, moths, and honeybees, sensory afferents from sensilla on the anterodorsal surface and the posteroventral surface of the flagellum form two nerves of almost equal thicknesses. In this study, double labeling of the two nerves, or proximal/distal regions of the nerves, with fluorescent dyes was used to investigate topographic organization of sensory afferents in the honeybee. The sensory neurons of ampullaceal sensilla responsive to CO₂, coelocapitular sensilla responsive to hygrosensory, and thermosensory stimuli and coeloconic sensilla of unknown function were characterized with large somata and supplied thick axons exclusively to the ventral nerve. Correspondingly, all glomeruli innervated by sensory tract (T) 4 received thick axonal processes exclusively from the ventral nerve. Almost all T1–3 glomeruli received a similar number of sensory afferents from the two nerves. In the macroglomerular complexes of the drone, termination fields of afferents from the two nerves almost completely overlapped; this differs from moths and cockroaches, which show heterogeneous terminations in the glomerular complex. In T1–3 glomeruli, sensory neurons originating from more distal flagellar segments tended to terminate within the inner regions of the cortical layer. These results suggest that some degree of somatotopic organization of sensory afferents exist in T1–3 glomeruli, and part of T4 glomeruli serve for processing of hygro‐ and thermosensory signals. J. Comp. Neurol. 515:161–180, 2009. © 2009 Wiley‐Liss, Inc.     

Inhibitory glutamate receptors in spider peripheral mechanosensory neurons

Most mechanosensory neurons are inhibited by GABAergic efferent neurons. This inhibition is often presynaptic and mediated by ionotropic GABA receptors at the axon terminals. GABA receptor activation opens Cl- channels, leading to membrane depolarization and an increase in membrane conductance. In many invertebrate preparations, efferent neurons that innervate mechanosensory afferents contain glutamate in addition to GABA, suggesting that the sensory neurons are also modulated by glutamate. However, the effects of glutamate on these neurons are not well understood. Peripheral parts of the spider (Cupiennius salei) mechanosensory neurons are surrounded by efferent fibers immunoreactive to antibodies against GABA and glutamate. GABA and its analogue muscimol were shown to effectively inhibit spider mechanosensory neurons innervating lyriform slit sensilla VS-3 that detects cuticular strains in the leg. Here, we show that glutamate also inhibits the VS-3 neurons, but its effects are different from those of GABA or muscimol, suggesting that it acts on a different group of receptors. GABA and muscimol always depolarized these neurons and the inhibitory effect was strongly correlated with the amount of depolarization. In contrast, glutamate inhibited the VS-3 neurons even when it did not depolarize them. In addition, while glutamate inhibited both the axonal action potentials elicited with electrical stimulation and dendritic action potentials produced by mechanical stimulation, muscimol only inhibited the axonal action potentials. Therefore, the inhibitory glutamate receptors in the VS-3 neurons are distinct from and differently distributed than the GABA receptors, providing a subtle control of the neurons' sensitivity in varying behavioural situations.     

Higher brain centers for social tasks in worker ants, Camponotus japonicus

Ants, eusocial insects, have highly elaborate chemical communication systems using a wide variety of pheromones. In the carpenter ant, Camponotus japonicus, workers and queens have the female-specific basiconic sensilla on antennae. The antennal lobe, the primary processing center, in female carpenter ants contains about 480 glomeruli, which are divided into seven groups (T1–T7 glomeruli) based on sensory afferent tracts. The axons of sensory neurons in basiconic sensilla are thought to project to female-specific T6 glomeruli. Therefore, these sensilla and glomeruli are thought to relate to female-specific social tasks in the ants. By using dye filling into local neurons (LNs) and projection neurons (PNs) in the antennal lobe, we neuroanatomically revealed the existence of an isolated processing system for signals probably relating to social tasks in the worker ant. In the antennal lobe, two categories of glomeruli, T6 glomeruli and non-T6 glomeruli, are clearly segregated by LNs. Furthermore, axon terminals of uniglomerular PNs from the respective categories of glomeruli (T6 uni-PNs and non-T6 uni-PNs) are also segregated in the secondary olfactory centers, the calyces of the mushroom body and the lateral horn: T6 uni-PNs terminate in the outer layers of the basal ring and lip of mushroom body calyces and in the posterior region of the lateral horn, whereas non-T6 uni-PNs terminate in the middle and inner layers of the basal ring and lip and in the anterior region of the lateral horn. These findings suggest that information probably relating to social tasks might be isolated from other olfactory information and processed in a separate subsystem.