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PubMed2002

Aberrant sensory innervation of the olfactory bulb in neuropilin-2 mutant mice.

The mammalian olfactory system consists of two anatomically segregated structures, the main olfactory system and the vomeronasal system, which each detect distinct types of chemical stimuli in the environment. During development, sensory neurons establish precise axonal connections with their respective targets within the olfactory bulb. The specificity of the odorant or vomeronasal receptor expressed by the sensory neuron is crucial in this process, yet it is less clear which of the more conventional axon guidance molecules are involved. Here, we show that neuropilin-2, a coreceptor for some of the class 3 semaphorins, is expressed in subpopulations of olfactory and vomeronasal sensory neurons. We generated a knock-out mutation in the neuropilin-2 gene by gene targeting in embryonic stem cells. Neuropilin-2 mutant mice exhibit profound and distinct effects on target innervation within the olfactory bulb. In the main olfactory system, axons of olfactory sensory neurons penetrate into the deeper layers of the main olfactory bulb. In the vomeronasal system, axonal fasciculation within the vomeronasal nerve is affected; some axons are misrouted and innervate glomeruli in an ectopic domain of the accessory olfactory bulb.

Animals↗
PubMed1976

Anesthetic-dependent field potential interactions in the olfactory bulb.

The effect of anesthetic upon olfactory bulb field potentials evoked by paired stimuli to the lateral olfactory tract (LOT) was examined in male rats. Experiments were performed on rats anesthetized with urethane or barbiturates and on unanesthetized rats with midcollicular brain stem sections. Anesthetic doses of urethane (1.5 g/kg) produced an increse in the test response for condition-test intervals of 15-60 msec or more. With barbiturate anesthesia there was a decrease in the test response for all intershock intervals up to 150 msec. In unanesthetized rats with midbrain sections there was an increase in the test response with low intensity shock pairs and a decrease in the test response with high intensity shock pairs. Simultaneous use of both anesthetics produced either an increase or a decrease in the test response, depending upon the relative doses and upon the stimulus intensity. Paired stimuli to the olfactory nerve produced results similar to those with LOT stimulation. Augmentation of the olfactory bulb field potential produced by paired olfactory nerve shocks was not abolished by lesions which sectioned the LOT and produced extensive damage of the anterior olfactory nucleus.

Animals↗
PubMed1986

Projection of septal organ receptor neurons to the main olfactory bulb in rats.

The septal organ of Masera is a distinct patch of olfactory epithelium on the septal wall of the nasal cavity. It lies ventral to the main olfactory receptor sheet. Central projections of the septal organ to the main and/or accessory olfactory bulb were studied with punctate HRP injections into the posterior dorsomedial olfactory bulb of rat pups. Injection sites encompassed 2-8% of the main olfactory bulb glomeruli and some injection sites were confined to the accessory olfactory bulb glomeruli. The distribution of receptor neurons in the septal organ that were labeled as a result of retrograde transport from the injection sites was determined. Olfactory receptor neurons were labeled in septal organs of those animals with injection sites confined to the main olfactory bulb. Subsequent to the posterior dorsomedial bulbar injections of this study, labeled receptor neurons were preferentially located in the posterior portion of the septal organ and were clustered about a plane dorsal to the plane bisecting the organ. Injections confined to the accessory olfactory bulb only labeled vomeronasal organ receptor neurons. Thus it is likely that the main olfactory bulb and not the accessory olfactory bulb is the bulbar terminus of septal organ receptor neurons. To date, experimental evidence suggests that the septal organ system may comprise an anatomically and functionally unique chemosensory pathway for odor detection.

Animals↗
PubMed1979

Effect of electrochemical stimulation in the olfactory bulbs on the release of gonadotropin hormones in rats.

The olfactory bulbs were electrochemically stimulated in non-anesthetized, freely moving rats, and the ovulatory response and serum luteinizing hormone (LH) levels were studied. The stimulus was applied at 11:30 h through unilateral stainless steel electrodes chronically implanted, and serial blood samples were obtained by way of a plastic cannula located in the jugular vein. Electrochemical stimulation (100 microA/60 sec) of the main olfactory bulbs in the ventral aspect of the superficial layers as well as in its central core, performed on the day of proestrus, prevented the preovulatory discharge of LH and ovulation. Blockade of ovulation also occurred when the stimulus was applied in the lateral or medial parts of the superficial layers. On the other hand stimulation in the dorsal part blocked the preovulatory surge of LH and ovulation in only 50% of the cases. Stimulation applied in the accessory olfactory bulbs failed to interfere with ovulation and LH discharge. Stimulation of ovariectomized estrogen-primed rats applied in superficial layers of the main olfactory bulbs had no significant effect on the release of LH, but when the stimulus was applied in the accessory olfactory bulb a release of LH took place 3-6 h following stimulation. These results indicate that stimulation of the olfactory bulbs produce a dual effect on LH release: while the response of the accessory olfactory bulb is facilitatory in nature, that of the main olfactory bulbs is inhibitory.

Animals↗
PubMed2004

Evidence for the disproportionate mapping of olfactory airspace onto the main olfactory bulb of the hamster.

Olfactory receptor neurons (ORNs) project to the rodent main olfactory bulb (MOB) from spatially distinct air channels in the olfactory recesses of the nose. The relatively smooth central channels of the dorsal meatus map onto the dorsal MOB, whereas the highly convoluted peripheral channels of the ethmoid turbinates project to the ventral MOB. Medial and lateral components of each projection stream innervate the medial and lateral MOB, respectively. To ascertain whether such topography entails the disproportionate representation seen in other sensory maps, we used disector-based stereological techniques in hamsters to estimate the number of ORNs associated with each channel in the nose and the number of their targets (glomeruli and mitral and tufted cells) in corresponding divisions of the MOB. Each circumferential half of the MOB (dorsal/ventral, medial/lateral) contained about 50% of the 3,100 glomeruli and about 50% of the 160,000 mitral and tufted cells per bulb. We found equivalent numbers of ORNs with dendritic knobs in the medial and lateral channels (4.5 million each). However, the central channels had only 2 million knobbed ORNs, whereas the peripheral channels had 7 million. Thus, there is a disproportionate mapping of the central-peripheral axis of olfactory airspace onto the dorsal-ventral axis of the MOB, encompassing a greater than threefold variation in the average convergence of ORNs onto MOB secondary neurons. We hypothesize that the disproportionate projections help to optimize chemospecific processing by compensating, with differing sensitivity, for significant variation in the distribution and concentration of odorant molecules along the olfactory air channels during sniffing.

Animals↗
PubMed2002

Grouping and representation of odorant receptors in domains of the olfactory bulb sensory map.

Individual glomeruli in the mammalian main olfactory bulb represent a single or at most a few types of odorant receptors. Thus the physical arrangement of glomeruli at the surface of the olfactory bulb can be viewed as a sensory map representing approximately 1,000 types of odorant receptors. This review summarizes the recent advance of the knowledge regarding the spatial organization of the sensory map in the main olfactory bulb. Recent studies show that individual olfactory bulbs contain dual sensory maps, one in the lateral hemisphere and the other in the medial hemisphere of the bulb. The tracings of selective subsets of olfactory axons to their target glomeruli in the olfactory bulb show that glomeruli are parceled into large zones or bands. The spatial arrangement of these zones and bands are stereotypical and conserved across individual mice. Optical imaging studies show that glomeruli in the most rostrodosal zone, zone I, are further parceled into smaller functional domains, and suggest that odorant receptors having a common or similar molecular feature receptive site are grouped together and represented by glomeruli within the functional domain. The possible relation between the functional domain organization and the subjectively perceived odor quality (olfactory submodality) is reviewed.

Axons↗
PubMed2000

Deafferentation-induced changes in the olfactory bulb of adult zebrafish.

The influence of the olfactory organ on maintenance of olfactory bulb structure was examined in zebrafish, using peripheral deafferentation. This fish provides a model in which the olfactory organ is easily accessible for removal, the animals easily survive the surgery, and the olfactory bulbs are small enough to allow rigorous analysis of the resulting effects. Unilateral olfactory organ ablations were performed on anesthetized adult zebrafish using a small-vessel cautery iron. Fish were allowed to survive for 1, 3, or 6 weeks following the procedure. Analysis of deafferented animals revealed that most, if not all, of the olfactory organ was missing on the ablated side, and the structure did not regenerate. The morphology of the olfactory bulb was affected notably by the removal of its primary afferent innervation. The olfactory nerve layer was diminished at 1 week and absent by 3 weeks post-deafferentation. At all of the survival times the deafferented bulb appeared significantly smaller at the gross level, and there was a statistically significant effect on bulb size and cell number after 6 weeks. Tyrosine hydroxylase expression, as revealed by immunohistochemistry, was decreased noticeably on the ablated side. In conclusion, the olfactory organ is important in the preservation of normal olfactory bulb anatomy and neurochemistry in adult zebrafish. Thus, the influence of the periphery does not end with the formation of the mature olfactory bulb.

Animals↗
PubMed1987

Olfactory bulb changes in Alzheimer's disease.

Olfactory bulbs in cases of Alzheimer's disease and age-matched controls have been examined by means of combining silver staining of pathological filaments with pigment-Nissl staining of the cell bodies. Neuritic plaques were found in the anterior olfactory nucleus. Neurofibrillary tangles and neuropil threads occurred in the anterior olfactory nucleus and in all layers of the olfactory bulb except the outer fibrous layer. The tangle-bearing neurons of the olfactory bulb were identified as tufted cells, outer granule cells, and two different types of nerve cells forming the rostral part of the anterior olfactory nucleus.

Alzheimer Disease↗
PubMed1994

Evidence for GABAB-mediated inhibition of transmission from the olfactory nerve to mitral cells in the rat olfactory bulb.

The GABAB agonist baclofen blocks transmission from the olfactory nerve to second order neurons in the frog olfactory bulb, and GABAB receptors in the rat olfactory bulb are selectively located in the glomerular layer. A reasonable hypothesis, therefore, is that inhibition in the glomerular layer is mediated, at least in part, by GABAB receptors. Here, we investigated the role of GABAB receptors in regulating the responses of mitral cells to activation of the olfactory nerve in the rat. Topical application of baclofen to the surface of the rat olfactory bulb reduced the amplitude of field potentials evoked by olfactory nerve stimulation (orthodromic response). Baclofen reduced the orthodromic response in a dose-dependent manner but the drug had no effect on the field potential evoked by antidromic activation of mitral cell axons (antidromic response). Baclofen also reduced olfactory nerve-evoked responses of mitral cells in an olfactory bulb slice preparation. The pharmacological specificity of the inhibition was confirmed by showing that the GABAB antagonist, CGP 55845A, blocked the inhibitory action of baclofen. These results suggest that transmission from olfactory nerve terminals to second order neurons is negatively regulated by periglomerular GABAergic interneurons; this inhibition is mediated, at least partially, by GABAB receptors.

Animals↗
PubMed1987

Locus coeruleus stimulation modulates olfactory bulb evoked potentials.

Field-evoked potentials from the main olfactory bulb in response to stimulation of the olfactory nerve and lateral olfactory tract were measured without and with conditioning stimulation of the locus coeruleus noradrenergic system. The locus coeruleus conditioning stimulus suppressed or inhibited the late components of the olfactory bulb potential evoked by orthodromic olfactory nerve stimulation; this inhibitory effect was suppressed by the microinjection of the alpha-adrenergic blocker prazosin into the olfactory bulb. Results indicate that noradrenergic fibers projecting from the locus coeruleus exert modulatory influences on neuronal networks underlying orthodromic evoked responses in the main olfactory bulb.

Adrenergic Fibers↗
PubMed1980

Subcellular distribution of glutamate decarboxylase in rat olfactory bulb: high content in dendrodendritic synaptosomes.

The olfactory bulbs in the CNS contain reciprocal dendrodendritic synapses between the granule cells and the secondary dendrites of mitral cells. Based on pharmacologic and electrophysiologic evidence, these synapses are believed to utilize GABA as an inhibitory neurotransmitter. A dendrodendritic synaptosomal fraction has been isolated from rat olfactory bulbs. The upper portion (PB) of the crude nuclear pellet contains 30-40% of the GAD (glutamate decarboxylase) activity of the olfactory bulb homogenate. When PB is purified on a discontinuous sucrose density gradient, 78-85% of the GAD activity is localized to the region containing the dendrodendritic synaptosomes, which were identified by transmission electron microscopy. The presence of a substantial proportion of GAD, the enzyme that catalyzes synthesis of GABA, in the DDS provides neurochemical support for the hypothesis that GABA functions at the reciprocal dendrodendritic synapses in the olfactory bulb.

Animals↗
PubMed1977

Absence of mitral cells in monolayer in monotremes. Variations in vertebrate olfactory bulbs.

An invariant feature of the olfactory bulb in placental and marsupial mammals is the arrangement of the perikarya of mitral cells in a monolayer. Contrasting with this is the arrangement found in the olfactory bulbs of the monotremes, platypus and echidna, where the large perikarya are not only absent from the position of a monolayer (usually forming the external boundary of the internal plexiform layer) but occupy a region which would characterize them as tufted cells. In other classes of amniote vertebrates, reptiles and birds, the placement of large perikarya in the olfactory bulb ranges from a compact layer to a broad band. Such an overview among several vertebrate classes suggests that a monolayer of mitral cells may be a specialized subset of the tufted-mitral cell population. The accessory olfactory formation among mammals also exhibits variation in the compactness of the large perikarya: a broadband in most but a compact layer in a few others such as the chinchilla and the capybara. Such specialized alignment of perikarya (and, consequently, of their dendritic and axonal elements) may enable more refined signal processing than does random alignment of these elements. Such speculations can be tested using appropriate phylogenetic sampling, and monotremes provide particularly advantageous test cases.

Anatomy, Comparative↗
PubMed1991

Olfactory bulb DA receptors may be located on terminals of the olfactory nerve.

The glomerular layer of the olfactory bulb contains a substantial population of dopaminergic neurons. We determined the quantity and location of D1 and D2 dopamine receptors which are the presumed targets of these neurons. Binding of the D1 selective ligand [3H]SCH23390 was slightly above background and was distributed through all layers of the bulb except the olfactory nerve layer. In contrast there were relatively high levels of [3H]spiperone binding to D2 DA receptors in the glomerular and olfactory nerve layers. The presence of relatively high concentrations of D2 DA receptors in both the nerve layer and glomerular layer suggests the novel hypothesis that these receptors may be localized on terminals of the olfactory nerve.

Animals↗
PubMed1999

Neural pathway from the olfactory bulbs regulating tonic gonadotropin secretion.

Removal of the olfactory bulbs of male golden hamsters results in a marked increase in tonic gonadotropin, prolactin and testosterone secretion which counteracts inhibitory effects of manipulations such as maintenance on short photoperiod, food restriction or treatment with gonadal steroids. The bulbectomy-induced increase in hormone secretion is interpreted to reflect a tonic inhibitory influence of the olfactory bulbs. This inhibition is not dependent upon chemosensory stimulation and may be mediated by olfactory bulb fibers projecting through the lateral olfactory tract to or through the olfactory tubercle. This review will relate these studies conducted on hamsters to results in other species, such as the rat, where the olfactory bulbs enhance serum gonadotropin levels.

Animals↗
PubMed1995

Olfactory experience modulates Bcl-2 expression in the developing olfactory bulb.

The effect of olfactory deprivation on cellular expression of the Bcl-2 gene in the olfactory bulb of young rats was investigated. Restriction of olfactory stimuli caused an overall increase in Bcl-2 mRNA expression, with increases seen in the lateral aspects of glomerular, external plexiform, mitral and granule cell layers, as well as the medial aspects of external plexiform layer. No differences were found in the unoperated control group. In addition, we found an inverse relationship between the incidence of apoptosis induced by olfactory deprivation and the magnitude of increase in Bcl-2 mRNA expression in the glomerular layer. These data raise the possibility that Bcl-2 may be involved in olfactory experience-related neural plasticity by regulating cell survival.

Animals↗
PubMed2003

Increase in the number of tyrosine hydroxylase-containing neurons in a primary culture system of the rat accessory olfactory bulb by co-culture with vomeronasal pockets.

Previously, we established a culture system of the accessory olfactory bulb in order to investigate the functional role of each accessory olfactory bulb neurons in pheromonal signal processing. In the present study, we developed a co-culture system of cultured accessory olfactory bulb neurons with partially dissociated cells of the vomeronasal organ. The dissociated cells of the vomeronasal organ form spherical structures surrounding a central cavity in culture, referred to as the vomeronasal pockets. The projection and activity of olfactory receptor neurons affect the differentiation and maturation of main olfactory bulb neurons. It was also reported induction of tyrosine hydroxylase expression in main olfactory bulb neurons when they were co-cultured with explants of the olfactory epithelium. Thus, we investigated the effects of co-culture with vomeronasal pockets on the differentiation and/or maturation of cultured accessory olfactory bulb neurons in relation to tyrosine hydroxylase expression. The number of tyrosine hydroxylase-containing neurons developmentally increased over time in the accessory olfactory bulb culture. This increase was significantly enhanced by coculture with vomeronasal pockets. Interestingly, a significant change in tyrosine hydroxylase expression was not observed when main olfactory bulb neurons were co-cultured with vomeronasal pockets. Moreover, significant changes in tyrosine hydroxylase expression were not observed when accessory olfactory bulb neurons were co-cultured with olfactory epithelium explants, as was previously observed in co-culture of main olfactory bulb neurons and olfactory epithelium explants. These results suggest that the differentiation and/or maturation of accessory olfactory bulb neurons is modified by vomeronasal organ neurons via specific interactions between the sensory organ and its target.

Animals↗
PubMed1997

Newly-generated cells from the rostral migratory stream in the accessory olfactory bulb of the adult rat.

Cell proliferation in the accessory olfactory bulb of the adult rat was analysed after systemic injection of 5-bromo-2'-deoxyuridine, detected immunocytochemically at different survival times and compared with proliferating cell nuclear antigen immunostaining. As previously described in the main olfactory bulb, local cell proliferation was absent or very limited. By contrast, starting from 15 days after bromodeoxyuridine administration, many immunoreactive nuclei were present in the granular layer, and to a lesser extent, in other layers of the accessory olfactory bulb. This suggests that the newly-generated cells are migrating elements of the rostral migratory stream which are known to reach the olfactory bulb in 15 days. By immunocytochemical detection of the polysialylated isoform of the neural cell adhesion molecule, a weakly-adhesive cell-surface molecule expressed by newly-generated/migrating cells of the rostral migratory stream, we found a high number of immunoreactive cells in the different layers of the accessory olfactory bulb. Most of these cells were observed in the granular layer and showed the morphology of migrating neuroblasts. Some immunoreactive cells displaying neuronal morphology were also detected in the external plexiform and glomerular layers. Double labelling experiments demonstrated that these cells are newly-generated cells. These results demonstrate the occurrence of newly-added cells in the accessory olfactory bulb of the adult rat, which likely correspond to the neuronal precursors originating from the rostral migratory stream. This could be relevant since the accessory olfactory bulb of rodents plays an important role in the hard wiring of a simple olfactory memory system for sexual pheromones.

Animals↗
PubMed1993

Immunohistochemical analyses of the human olfactory bulb.

These studies explore the distribution of putative neuroactive peptides in the human olfactory bulb. Localization of synaptophysin-, serotonin-, cholecystokinin-, substance P-, and somatostatin-like staining was examined by immunocytochemical protocols. The results provide new insights into the composition and laminar segregation of subpopulations of neurons and neuronal processes in the human olfactory bulb. The prominent synaptophysin-like immunoreactivity observed in the glomeruli of the human olfactory bulb is consistent with the notion that the density of synapses, and hence the density of synaptic vesicles, is highest in the glomeruli. Serotonin-like immunoreactivity suggested a variable innervation of glomeruli ranging from a dense tangled ball of fibers within the glomerulus to a sparse innervation by a single immunoreactive fiber. There was no evidence of serotonin-like immunoreactive cell bodies in either the olfactory bulb proper, anterior olfactory nucleus, or proximal regions of the lateral olfactory tract. Cholecystokinin-like immunoreactivity was limited to fibers found largely in the juxtaglomerular region of the glomerular layer. In the deeper layers of the olfactory bulb, cholecystokinin-like immunoreactive fibers did not show any of branching or arborization that was evident in the juxtaglomerular region. Substance P-like immunoreactivity was seen in varicose fibers distributed in all of the human olfactory bulb laminae. In addition, stained multipolar neurons were found in the area of the anterior olfactory nucleus. Somatostatin-like immunoreactivity was similar to that of substance P in that a plexus of stained fibers was found in all laminae of the olfactory bulb. Also, somatostatin-like immunoreactive cell bodies were found in the area of the anterior olfactory nucleus. However, as compared to substance P, somatostatin had a less dense plexus of immunoreactive fibers in the olfactory bulb. These results increase our understanding of the fundamental organization of the human olfactory system. The current data, coupled with prior studies, provide a foundation from which to study the cellular pathology of diseases with known olfactory system sequelae such as Alzheimer's, Parkinson's, and schizophrenia.

Adult↗