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PubMed2006

Female snake sex pheromone induces membrane responses in vomeronasal sensory neurons of male snakes.

The vomeronasal organ (VNO) is important for activating accessory olfactory pathways that are involved in sexually dimorphic mating behavior. The VNO of male garter snakes is critically important for detection of, and response to, female sex pheromones. In the present study, under voltage-clamp conditions, male snake VNO neurons were stimulated with female sexual attractiveness pheromone. Thirty-nine of 139 neurons exhibited inward current responses (reversal potential: -10.6 +/- 2.8 mV). The amplitude of the inward current was dose dependent, and the relationship could be fitted by the Hill equation. Under current-clamp conditions, application of pheromone produced membrane depolarizing responses and increases in firing frequency. These results suggest that the female pheromone directly affects male snake VNO neurons and results in opening of ion channels, thereby converting the pheromone signal to an electrical signal. The response to female pheromone is sexually dimorphic, that is, the pheromone does not evoke responses in VNO neurons of female snakes. An associated finding of the present study is that the female sex pheromone, which is insoluble in aqueous solutions, became soluble in the presence of Harderian gland homogenate.

Animals↗
PubMed1993

GABAergic control of odor-induced activity in the frog olfactory bulb: electrophysiological study with picrotoxin and bicuculline.

In the olfactory bulb, the first relay of the olfactory pathways, GABA, could be largely involved in the information processing since the two main populations of interneurons, periglomerular and granular cells, use it as neurotransmitter through reciprocal synapses with second-order neurons. This study planned to clarify the role of GABAergic inhibition in odor coding and, more precisely, the role of glomerular GABAergic inhibition. To do so, we attempted to specifically block in vivo GABAA receptors with either picrotoxin or bicuculline. The drug was applied at the level of the glomerular layer so that the antagonist could act primarily via periglomerular cells. The analysis of the effects of blocking GABAA on the coding was studied by recording the second-order neuron responses to odor stimuli delivered in a wide concentration range. Under drug treatment, the second-order neuron properties were deeply changed: response thresholds to odors were often lowered and spike bursts were more sustained in frequency and in duration. Thus, the GABAergic control on second-order neurons might be carried out by limiting the neuron excitability. GABAA antagonists applied in this manner could act to suppress the inhibitory effect of either the periglomerular cells or of the granule cells, both of which have been shown to contain enzymes for GABA production. The placement of the drug suggests to us that the action is primarily at the glomerulus. The results are consistent with periglomerular cells exerting a tonic inhibition on second-order neurons, an inhibition whose strength would be modulated by stimulus intensity. As a result, the amplifying role of glomerular convergence might be partly counterbalanced by input inhibition. Nevertheless, due to our procedure of drug application, one cannot rule out the possibility that the effects observed may partly reflect granular cell blocking. It can be concluded that the whole GABAergic inhibition, through GABAA receptors, permits a wide dynamic range of intensity coding.

Action Potentials↗
PubMed1995

Odor responses after complete desensitization of the cAMP-dependent pathway in turtle olfactory cells.

The degree of contribution of the cAMP signal transduction pathway to odor responses was examined by recording current responses from isolated turtle olfactory cells under the whole-cell voltage clamp conditions. The cAMP signal transduction pathway was desensitized by dialyzing 1 mM cAMP and 0.5 mM IBMX from the patch pipette into the cells. Extracellular application of 3 mM cpt-cAMP, a membrane-permeable cAMP analogue elicited no response, indicating that the cAMP pathway was completely desensitized. Application of an odorant cocktail induced a large inward current under these conditions, suggesting that the cAMP-independent signal transduction contributes significantly to generation of odor responses in the turtle.

1-Methyl-3-isobutylxanthine↗
PubMed1996

Development of olfactory glomeruli: temporal and spatial interactions between olfactory receptor axons and mitral cells in opossums and rats.

Mitral cells are the primary output neurons of the vertebrate olfactory bulb and are major recipients of sensory input from the periphery. The morphogenesis of mitral cell dendrites was followed to elucidate their early spatial and temporal interactions with olfactory receptor neurons and glia during the construction of olfactory glomeruli. Monodelphis domestica, a marsupial born at an extremely immature stage, and rats were examined. Mitral cells were retrogradely labeled by application of the lipophilic dye 1,1' dihexadecyl-3,3,3'3'-tetramethylin-docarbocyanine perchlorate (DiI) to the lateral olfactory tract. In double-labeling experiments, olfactory receptor neurons were stained with 3,3' dihexadecyloxacarbocyanine perchlorate (DiO), or olfactory nerve Schwann cells were visualized using S-100 protein immunohistochemistry. Tissue was examined with a confocal laser scanning microscope. Some preparations were subsequently investigated with an electron microscope. In Monodelphis, differentiation of mitral cells starts with an outgrowth of numerous, uniform, and widespread dendrites. As soon as terminals of olfactory receptor axons coalesce into glomerular knots within the presumptive glomerular layer, dendrites of individual mitral cells innervate several adjacent glomeruli where they receive sensory synaptic input. With maturation, supernumerary mitral cell dendrites retract, leaving one primary dendrite bearing a terminal glomerular tuft. Simultaneously, secondary dendrites begin to arise. The formation of glomeruli begins earlier and progresses faster in the rat compared to Monodelphis. Nevertheless, mitral cell differentiation in both species follows a common sequence: overproduction of dendrites, selection of usually one primary apical dendrite, and elimination of supernumerary processes. Since olfactory receptor neurons form synaptic contacts with the widespread mitral cell dendrites, considerable synaptic rearrangement must occur within the olfactory glomeruli during maturation.

Animals↗
PubMed1996

Glial cells stabilize axonal protoglomeruli in the developing olfactory lobe of the moth Manduca sexta.

Odor information is processed in spherical structures called glomeruli, which in all animals with differentiated olfactory systems are sites of densely spaced synaptic contacts between olfactory sensory axons and target central nervous system (CNS) neurons. Glomerulus development in the antennal (olfactory) lobe of the moth brain, which is initiated by the arrival of antennal receptor axons, requires interaction among three elements: glial cells, receptor axons, and their targets, the antennal-lobe neurons. Receptor axons form an array of protoglomeruli that become surrounded by glia and serve as a template for mature glomeruli. Previous experiments showed that when the number of glial cells is sharply reduced during development either by irradiation or by an anti-mitotic agent, receptor axons form protoglomeruli, but in the mature lobes, glomeruli are absent and central neurons lack the characteristic glomerular tufting of their arbors. The current investigation was conducted to determine which cellular events in the process of glomerulus formation are disrupted by severe reduction in glial-cell number. The branching patterns of receptor axons and antennal-lobe neurons were examined in animals that had been irradiated to produce glia-deficient antennal lobes at stages during which glomeruli normally would develop. We found that the receptor axons did form protoglomeruli, but that the protoglomeruli quickly disintegrated in glia-deficient antennal lobes; the receptor axons branched diffusely, except where several neighboring glia survived irradiation and together formed a wall of processes that appeared to block the passage of neuronal processes. Multi-glomerular antennal-lobe neurons never developed tufted arbors even at early stages. These results suggest that maintenance of protoglomeruli depends on the border of glia that forms around each protoglomerulus and that the subsequent tufting of antennal-lobe neurons depends on maintenance of the protoglomerular template during the period of dendritic growth.

Animals↗
PubMed1997

Differential expression of manganese and copper-zinc superoxide dismutases in the olfactory and vomeronasal receptor neurons of rats during ontogeny.

Superoxide dismutases (SODs) protect cells from damage by oxygen free radicals. Manganese (Mn) SOD is preferentially induced in terminally differentiating cells; induction of copper-zinc (CuZn) SOD is more closely associated with postnatal exposure to environmental sources of oxygen free radicals. The purpose of this study was to investigate ontogenetic changes in immunoreactivity for MnSOD and CuZnSOD relative to the expression of markers of neuronal and chemosensory differentiation in olfactory and vomeronasal receptor neurons (ORNs and VRNs, respectively), which mature with different time courses. Immunoreactivity for both SODs was detected in rat ORNs at embryonic day (E) 14, the earliest time point investigated, but not until E16 in vomeronasal neuroblasts. ORNs also expressed the neuronal marker protein gene product (PGP) 9.5 and the chemosensory cell marker olfactory marker protein (OMP) at E14; vomeronasal neuroblasts expressed PGP 9.5 at E16 but were not immunoreactive for OMP until postnatal day (P) 2. Immunoreactivity for MnSOD in ORNs and VRNs generally increased pre- and postnatally to a maximum at P11. Immunoreactivity for CuZnSOD did not increase markedly until after birth, reaching maximal levels at P11-P24. Within ORNs and VRNs, the most intense immunoreactivity was localized in the dendritic and supranuclear regions. The results indicate that in ORNs and VRNs, increases in MnSOD immunoreactivity during ontogeny parallel the ongoing differentiation and maturation of chemosensory receptor neurons; in contrast, the induction of immunoreactivity for CuZnSOD is associated with postnatal exposure to the ambient oxygen and xenobiotic environment.

Age Factors↗
PubMed1998

Differential projections of ciliated and microvillous olfactory receptor cells in the catfish, Ictalurus punctatus.

The primary olfactory projections of channel catfish Ictalurus punctatus have been examined with postmortem tracing by using either 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate or 1,1-dilinoleyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate (DiI). Following DiI deposition into small areas in different parts of the posterior olfactory bulb, olfactory sensory neurons always were labeled throughout the olfactory epithelium. No obvious topographical mapping exists between the epithelium and olfactory bulb. The different dye placements, however, did result in labeling of different morphologies of receptor cells, depending on the site of injection. Retrogradely labeled neurons in the olfactory epithelium were classified into three types on the basis of their height: tall, intermediate, and short. Tall olfactory sensory neurons had perikarya at the bottom one-fourth of the epithelium, extended slender dendrites to the epithelial surface, and possessed numerous cilia on the apical dendritic tips. These tall olfactory sensory neurons were labeled predominantly following DiI applications to the ventral part of the posterior olfactory bulb. In contrast, the short olfactory sensory neurons had perikarya situated within the superficial half of the epithelium and with short apical dendrites bearing microvilli. These short olfactory sensory neurons projected predominantly to the dorsal, posterior olfactory bulb. Thus, short microvillous receptor cells and tall ciliated receptor cells connect to different parts of the olfactory bulb, although the receptor cells are intermingled within the olfactory epithelium. Because different parts of the olfactory bulb are thought to respond preferentially to different classes of odorants, these results suggest that receptor cell morphology may be related to odorant quality detection. In addition, to compare this study with previous in vivo studies, Fluoro-Gold was injected in vivo into either the olfactory bulb or intraperitoneally. These in vivo studies show that so-called "type II ciliar receptor cells" of the nonsensory epithelium are labeled nonselectively by blood-borne substances, but they are not labeled by postmortem injections of DiI anywhere in the olfactory bulb.

Animals↗
PubMed1998

Neurogenesis and migration of receptor neurons in the vomeronasal sensory epithelium in the opossum, Monodelphis domestica.

The sensory epithelium of the vomeronasal organ (VNO) contains primary chemosensory receptor neurons that project to the accessory olfactory bulb (AOB). In the present study, neurogenesis and cell migration in the sensory epithelium of the VNO were analyzed in opossums (Monodelphis domestica) by using bromodeoxyuridine (BrdU) labeling. 1) In the VNO of normal adult opossums, BrdU labeled a small number of cells localized in the basal region of the sensory epithelium. After 1 or 2 weeks of survival, the labeled cells appeared in the receptor cell layers and became receptor neurons, as indicated by coexpression of the G proteins G(i alpha2) or G(o alpha). 2) In the VNO in which the receptor neurons had been destroyed by removing the AOB, the number of BrdU-labeled cells in the reconstituting sensory epithelium was greatly increased compared with that in the intact VNO. The labeled cells were also located in the basal region of the sensory epithelium. 3) In the developing VNO (at postnatal day 10), more cells in the basal region of the sensory epithelium were labeled than in the adult VNO, indicating rapid cell proliferation; and there appeared to be more labeled cells in the basal region near the margins of the sensory epithelium where it meets the nonsensory epithelium. These observations demonstrate that, in the opossum VNO, there is a population of proliferating cells in the basal region close to the basal lamina in the sensory epithelium. The newly generated neurons in the basal region migrate vertically into the receptor cell layer.

Animals↗
PubMed1999

Lobster G-protein coupled receptor kinase that associates with membranes and G(beta) in response to odorants and neurotransmitters.

A cDNA clone (lobGRK2) encoding a protein of 690 amino acids with significant similarity to the GRK2 subfamily of G-protein coupled receptor kinases was isolated. lobGRK2 was widely expressed as a 9-kb major transcript and a protein of 80 kDa. It was most abundant in the brain and the olfactory organ but was absent in the eye/eyestalk. Immunocytochemistry revealed lobGRK2 immunoreactivity in the outer dendritic segments of the olfactory receptor neurons, the site of olfactory transduction. LobGRK2 immunoreactivity was observed in most neuronal structures in the brain, although with varying intensity. It was strongest in neuropil, especially the olfactory and accessory lobes but was also detectable in neuronal cell bodies. Stimulation of brain homogenates with a mixture of neurotransmitters increased the association of lobGRK2 with membranes and with G(beta). Similarly, stimulation of olfactory dendrite homogenates with an odorant mixture caused lobGRK2 to associate with G(beta). These results support the conclusion that lobGRK2 responds to odorants and to neurotransmitters and may act to initiate desensitization by phosphorylating G-protein-coupled receptors in the olfactory organ and the brain, respectively.

Animals↗
PubMed2000

Pattern of calretinin immunoreactivity in the main olfactory system and the vomeronasal system of the tree shrew, Tupaia belangeri.

The distribution of the calcium-binding protein calretinin was studied in peripheral and central parts of the main olfactory system (MOS) and the vomeronasal system (VNS) of adult tree shrew Tupaia belangeri. The calretinin immunoreaction was carried out with a peroxidase-coupled polyclonal antibody. In the VNS, complete labeling of all receptor cells and vomeronasal nerve fibers was observed, whereas only a subset of the somata and dendrites of receptor cells and of the olfactory nerve fibers of the MOS was immunoreactive. From the immunoreactive dendritic clubs of vomeronasal receptor cells, calretinin-labeled structures, presumably clumps of microvilli, arose that terminated within immunopositive portions of the mucus. In the main olfactory bulb, the neuropil of some of the glomeruli was immunoreactive. All periglomerular and many mitral cells were labeled. The external plexiform layer was subdivided into a faintly immunoreactive superficial half and a strongly immunoreactive deep half. Immunoreactive basal dendrites of mitral cells could be followed into either the deep half or the superficial half. In the laminated internal granular layer, a subset of immunopositive granule cells extended dendrites into the external plexiform layer. Mitral cells and granule cells with dendrites ascending to different levels of the external plexiform layer may represent functional subclasses. In the accessory olfactory bulb, all vomeronasal nerve fibers, glomeruli, and mitral/tufted cells were labeled, whereas immunoreactive periglomerular cells and internal granule cells were only scattered. In Tupaia, calretinin immunoreactivity is a more general property of the primary projecting neurons of the VNS than of the MOS and possibly indicates the involvement of calretinin in the perception of certain of the olfactory qualities.

Animals↗
PubMed1996

Information processing in mammalian olfactory system.

In recent years, considerable progress has been made in understanding how the olfactory system uses neural space to encode sensory information. In this review, we focus on recent studies aimed at understanding the organizational strategies used by the mammalian olfactory system to encode information. The odorant receptor gene family is discussed in the context of its genomic organization as well as the specificity of olfactory sensory neurons. These data have important consequences for the mechanisms of odorant receptor gene choice by a given sensory neuron. Division of the olfactory epithelium into zones that express different sets of odorant receptors is the first level of input organization. The topographical relationship between periphery and olfactory bulb represents a further level of processing of information and results in the formation of a highly organized spatial map of information in the olfactory bulb. There, local circuitry refines the sensory input through various lateral interactions. Finally, the factors that may drive the development of such a spatial map are discussed. The onset of expression and the establishment of the zonal organization of odorant receptor genes in the epithelium are not dependent upon the presence of the olfactory bulb, suggesting that the functional identity of olfactory sensory neurons is determined independently of target selection.

Animals↗
PubMed1996

Perireceptor events in olfaction.

Before reaching olfactory receptor neurons, odorant molecules have to cross an aqueous interface: the nasal mucus in vertebrates and the sensillar lymph in insects. Biochemical interactions taking place between odorants and the elements of these phases are called perireceptor events. Main protein constituents of these media, in both insects and vertebrates, are OBPs (odorant-binding proteins). Another class of proteins active in the olfactory perireceptor area includes odorant-degrading enzymes. The structure and the properties of these major proteins, with particular reference to OBPs, are reviewed and their role in olfactory transduction is discussed.

Amino Acid Sequence↗
PubMed1999

Calcium-binding proteins: differential expression in the rat olfactory cortex after neonatal olfactory bulbectomy.

Calbindin, parvalbumin, and calretinin, members of EF-hand calcium-binding proteins, play important roles in buffering intracellular calcium ions. These proteins are localized in distinct populations of cells in the olfactory bulb (the primary sensory relay in the olfactory system) and its major synaptic target, the primary olfactory cortex (POC). In the present study, the postnatal expression of these calcium-binding proteins in layer III of POC was quantitatively examined 30 days after neonatal bulbectomy, a manipulation known to cause cell death and neurotransmitter changes. The numbers of both calbindin and parvalbumin-immunoreactive profiles showed significant increases (68% and 163%, respectively), while calretinin-immunoreactive profiles exhibited a 46% reduction. The data demonstrate that the expression of these calcium-binding proteins is regulated in part by the afferent input from the olfactory bulb. Furthermore, the resultant increase in calbindin and parvalbumin expression may provide neuroprotective support necessitated by possible alterations in intracellular calcium ions and other neurochemical factors that accompany neonatal bulb removal.

Animals↗
PubMed1999

Widespread expression of olfactory cyclic nucleotide-gated channel genes in rat brain: implications for neuronal signalling.

The cyclic nucleotides cAMP and cGMP are important intracellular messengers involved in a wide variety of signal transduction events in the nervous system. It has been proposed that cAMP/cGMP elicit some of their effects through direct gating of a novel class of Ca2+ -permeable ion channels that are termed cyclic nucleotide-gated (CNG) channels. Previous studies have identified the expression of a gene encoding one major CNG channel subtype, the olfactory receptor neuron alpha subunit, in the brain [El-Husseini et al. (1995) NeuroReport 6:1331-1335; Kingston et al. (1996a) Proc. Natl. Acad. Sci. U.S.A. 93:10440-10445; Bradley et al. (1997) J. Neurosci. 17:1993-2085]. We, therefore, proposed that the actions of cAMP/cGMP on neurons in the brain might occur through the activation of these CNG channels. To determine how widespread such a function might be, the regional and cellular distribution of the olfactory CNG channel alpha subunit has been examined in detail. Primers for multiple portions of the olfactory CNG channel were used in polymerase chain reaction (PCR) to amplify cDNA reverse-transcribed from several brain regions. The identities of PCR products were confirmed with Southern blots and by sequencing. In situ hybridization experiments demonstrated localization of CNG channel mRNA in discrete neuronal populations throughout the brain. In agreement with previous work, relatively strong hybridization signals are present in neuronal cell bodies of the cerebellum, olfactory bulb, cerebral cortex, and brainstem. Additionally, somewhat lesser signals are found in thalamus, hypothalamus, midbrain, and spinal cord while no hybridization signal was detectable in the caudate nucleus. This surprisingly wide distribution throughout the rat brain strengthens the hypothesis that CNG channels may influence numerous processes as downstream effectors of cyclic nucleotide cascades. Interestingly, the distribution of CNG channels is very similar to that of the nitric oxide/cGMP system, suggesting that one function of CNG channels in the brain could be to link diffusible messengers to elevated Ca2+ entry into neurons.

Animals↗
PubMed2000

Patterning of olfactory sensory connections is mediated by extracellular matrix proteins in the nerve layer of the olfactory bulb.

In early rat embryos when axons from sensory neurons first contact the olfactory bulb primordium, lactosamine-containing glycans (LCG) are detected on neurons that are broadly distributed within the olfactory epithelium, but that project axons to a very restricted region of the ventromedial olfactory bulb. LCG(+) axons extend through channels defined by the coexpression of galectin-1 and beta2-laminin. These two extracellular matrix molecules are differentially expressed, along with semaphorin 3A, by subsets of ensheathing cells in the ventral nerve layer of the olfactory bulb. The overlapping expression of these molecules creates an axon-sorting domain that is capable of promoting and repelling subsets of olfactory axons. Specifically, LCG(+) axons preferentially grow into the region of the nerve layer that expresses high amounts of galectin-1, beta2-laminin, and semaphorin 3A, whereas neuropilin-1(+) axons grow in a complementary pattern, avoiding the ventral nerve layer and projecting medially and laterally. These studies suggest that initial patterning of olfactory epithelium to olfactory bulb connections is, in part, dependent on extracellular components of the embryonic nerve layer that mediate convergence and divergence of specific axon subsets.

Age Factors↗
PubMed1993

The cellular and genetic basis of olfactory responses in Caenorhabditis elegans.

The small soil nematode Caenorhabditis elegans has only 302 neurons in its entire nervous system, so it is possible to analyse the functions of individual neurons in the animal's behaviour. We are using behavioural, cellular and genetic analyses of chemotactic responses to find out how olfactory behaviour patterns are generated and regulated. Single chemosensory neurons in C. elegans can recognize several different attractive odorants that are distinguished by the animal. Distinct sets of chemosensory neurons detect high and low concentrations of a single odorant. Odorant responses adapt after prolonged exposure to an odorant; this adaptation is odorant specific and reversible. Mutants with defects in odorant responses have been identified. Some genes appear to be necessary for the development or function of particular kinds of sensory neurons. Other genes have effects that suggest that they participate in odorant reception or signal transduction.

Adaptation, Physiological↗
PubMed1993

Membrane currents and mechanisms of olfactory transduction.

The term olfactory transduction refers to the mechanisms that transform chemical information into electrical signals. With the patch-clamp technique it is possible to record those signals and to infer something about the mechanism that produced them. The direct activation of a cation-permeable channel by cAMP is the final step in producing the odour-induced ionic current. Because it occupies a critical position in the transduction process, measurements of the ion channel's activity provide useful insights into the molecular processes underlying olfactory transduction. In addition to its activation by cAMP and cGMP, the channel is modulated by both extracellular and intracellular Ca2+ ions and by extracellular Mg2+ ions, all at physiological concentrations. These effects are probably important in promoting signal reliability. An unusual feature of this channel is its termination kinetics--it can remain active for hundreds of milliseconds after the agonist has been removed. This is likely to add to the integrating properties of the olfactory sensory neuron.

Animals↗
PubMed1975

A degeneration study of some habenular efferents to the midbrain in a wallaby.

In this study, specimens of both sexes of a Western Australian wallaby (Steonix brachyurus) had surgical or electrolytic lesions made in the habenular complex unilaterally. Thirty-five-micron sections, cut coronally and parasagittally, were stained by the Nauta-Gygax ('54) or the Fink-Heimer ('67) technique to demonstrate degenerating fibers. Degeneration was seen in the habenular commissure and bilaterally in the habenula and fasciculus retroflexus, but was most extensive ipsilaterally. Habenular fibers destined for the tegmentum and central gray passed caudally into the midbrain in a continuous fan-like array which extended from the central gray dorsally to the interpeduncular nucleus ventrally. In their posterior course some fibers crossed the midline with the decussating cerebellar fibers. The majority of fibers, with those in the fasciculus retroflexus which passed ventral to the decussation of the superior cerebellar peduncle, turned dorsally. They passed mostly just lateral to the midline nuclei and into the central gray where degeneration was seen throughout the entire extent. Terminal degeneration was seen in the interpeduncular nucleus primarily in the dorsal and posterior parts, in the ventral tegmental nucleus and throughout the central gray, but was more densely clustered in the region of the dorsal tegmental nucleus.

Animals↗