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PubMed1991

Differential projections of the main and accessory olfactory bulb in the frog.

The central projections of the main olfactory bulb and the accessory olfactory bulb of the adult leopard frog (Rana pipiens) were reexamined, by using a horseradish peroxidase anterograde tracing method that fills axons with a continuous deposit of reaction product. The fine morphology preserved by this method allowed the terminal fields of the projection tracts to be delineated reliably, and for the first time. Herrick's amygdala has been newly subdivided into cortical and medial nuclei on the basis of cytoarchitecture, dendritic morphology, and the differential projections of the main and accessory olfactory tracts. The main olfactory bulb projects through the medial and lateral olfactory tracts to the postolfactory eminence, the rostral end of the medial cortex, the rostral end of the medial septal nucleus, the cortical amygdaloid nucleus, the nucleus of the hemispheric sulcus, and both the dorsal and ventral divisions of the lateral cortex, including its retrobulbar fringe. The lateral olfactory tract overlaps the dorsal edge of the striatal plate along the ventral border of the lateral cortex, but it is not certain whether any striatal cells are postsynaptic to the tract fibers. The lateral cortex is the largest of these territories, and receives the terminals of the main olfactory projection throughout its extent. It extends from the olfactory bulb to the posterior pole, and from the striatum to the summit of the hemisphere, where it borders the dorsal cortex. The medial and lateral olfactory tracts combine in the region of the amygdala to form a part of the stria medullaris thalami. These fibers cross in the habenular commissure and terminate in the contralateral cortical amygdaloid nucleus and periamygdaloid part of the lateral cortex. Cells projecting to the main olfactory bulb are found in the diagonal band and adjacent cell groups, but there is no evidence of an interbulbar projection arising from either the olfactory bulb proper or a putative anterior olfactory nucleus. The accessory olfactory bulb projects through the accessory olfactory tract to the medial and cortical amygdaloid nuclei. A fascicle of the tract crosses in the anterior commissure to terminate in the contralateral amygdala. While the main and accessory olfactory projections may converge in the cortical amygdaloid nucleus, the medial amygdaloid nucleus is connected exclusively with the accessory olfactory bulb.

Afferent Pathways↗
PubMed1994

The development of glial fibrillary acidic protein-positive cells and the appearance of laminin-like immunoreactivity in fetal olfactory bulb transplants.

Embryonic rat olfactory bulbs were transplanted into the site vacated by aspiration of an olfactory bulb from a neonatal rat. This paper presents our findings related to the development of glial fibrillary acidic protein (GFAP) positive (+ve) glial cells and the appearance of laminin-like immunoreactivity in these transplants. The GFAP+ve glial cells formed perivascular end-feet on the invading vasculature and formed a glia limitans along the pial surface of the transplant. This reconstituted glia limitans was continuous with that of the host brain, there being no glia limitans at the donor-host interface. Thus, the donor tissue was well-integrated with that of the host brain.

Animals↗
PubMed1990

Modification of olfactory bulb synaptic inhibition by early unilateral olfactory deprivation.

Early unilateral olfactory deprivation produces large structural and neurochemical changes in the olfactory bulb, the first central relay for olfactory information. The functioning of deprived bulbs was examined in the present report by using paired-pulse stimulation of the lateral olfactory tract. Paired-pulse stimulation reflects interactions between mitral/tufted cells and granule cells, as well as the modulatory effects of centrifugal and intra-bulbar association fibers. Paired-pulse stimulation produced inhibition of mitral/tufted cells in control animals at PN20-PN22. This inhibition was significantly enhanced in littermates deprived of olfactory input from PN1 to PN20-PN22. Suppression of mitral/tufted cell single-unit spontaneous activity following single-pulse stimulation of the lateral olfactory tract (LOT) was similarly enhanced in deprived bulbs. These results suggest that early olfactory deprivation significantly modifies subsequent olfactory system function.

Animals↗
PubMed1992

gamma-Aminobutyric acid activity in the olfactory bulb of the rat during the sexual cycle and response to olfactory stimuli.

Glutamic acid decarboxylase activity in the main and accessory olfactory bulbs throughout the sexual cycle of the rat was studied. The effect of male pheromonal secretion on enzyme activity during proestrus and estrus day was also tested. The enzyme activity showed circadian rhythm during the estrous cycle. This rhythm was disrupted during diestrus-2 afternoon in the main bulb and came back during proestrus afternoon. A different pattern of enzyme activity was present in the accessory bulb, since the circadian rhythm was altered during proestrus morning, returning during estrus afternoon. Male odor exposition did not change enzyme profile activity during proestrus day and during estrus morning in the main bulb. In contrast, in the accessory bulb the olfactory stimuli induced opposite changes to that found in rats from the vivarium during proestrus. Comparison of enzyme activity in olfactory stimuli-deprived rats with that of pheromone-stimulated rats during proestrus showed that male odor exposure specifically affects accessory bulb enzyme activity. It is concluded that the changes of the olfactory bulb GABAergic system during proestrus and estrus day, or that evoked by odor stimuli, demonstrate the discriminative response of this system between the accessory olfactory bulb and the main olfactory bulb.

Animals↗
PubMed1993

Terminal arborizations of olfactory nerve fibers in the glomeruli of the olfactory bulb.

The glomerulus of the olfactory bulb may serve as a fundamental organizational unit for odor representation. In this context, the axons of olfactory receptor cells with similar response spectra may converge in specific glomeruli. While the topography between the olfactory epithelium and the olfactory bulb glomeruli has been explored, the characteristics of primary afferent terminal fields within glomeruli are poorly understood. To explore this issue, reconstructions of the terminal arbors of single olfactory nerve (ON) fibers within glomeruli were carried out in the rat olfactory bulb at the light microscopic level. Tissue samples prepared with the Golgi-EM technique resulted in distinct impregnation of limited subsets of individual ON fibers. Following camera lucida reconstruction, quantitative analyses were made on selected sets of ON fibers and on the glomeruli they invaded. Most ON fibers began to arborize only after penetrating the glomeruli to a mean depth of approximately 35.9 microns. The fibers gave rise to a complex arbor of branches that was limited in total length (mean 157.7 microns) and the number of en passant varicosities or terminal enlargements (mean = 8.1). The number of varicosities and terminal boutons was proportional to the total length of branches. Also, there was a statistically significant correlation between the total branch length and the area (microns2) encompassed by the total arbor. The intraglomerular region supplied by the terminal arbor of an individual ON fiber was generally restricted. Given the recent molecular specificity attributed to olfactory receptor cells, this may provide a morphological basis for selective processing of signals encoded by specific receptors in the olfactory epithelium.

Animals↗
PubMed1976

[Electrotonic interaction of secondary neurons of the carp olfactory bulb].

Responses of the olfactory bulb secondary neurons provoked by olfactory tract electrical stimulation were studied in carp, using the intracellular recording technique. Spike responses in most neurons were identified as antidromic. Values of their latent periods ranged from 2.5 to 55 msec. In addition, two kinds of responses with a constant latent period were revealed: pseudoantidromic spikes and low-amplitude fast depolarizing potentials. It is supposed that both kinds of responses are generated by an antidromic spike of the neighbouring secondary neuron electrotonically coupled with the neuron recorded.

Action Potentials↗
PubMed1988

Spatial patterns of olfactory bulb single-unit responses to learned olfactory cues in young rats.

1. Neonatal rat pups were classically conditioned to an odor stimulus from postnatal day 1 (PN1) to PN18. Tactile stimulation (stroking) was used as the unconditioned stimulus. On PN19, mitral/tufted cell single-unit responses to the conditioned odor were examined in both conditioned and control pups. Recordings were made from mitral/tufted cells in two regions of the olfactory bulb: 1) an area typically associated with focal [14C]2-deoxyglucose (2-DG) uptake in response to the conditioned odor and 2) an area distant from focal 2-DG uptake to the conditioned odor. Animals were anesthetized with urethane and were naturally respiring during the single-unit recording procedure. 2. Changes in mitral/tufted cell firing rate in response to odors in both bulbar regions and all training groups were classified as either excitatory, suppressive, or no response. This response classification was used to compare response patterns to the conditioned odor between bulbar regions and training groups. 3. Classical conditioning selectively modified the response patterns of mitral/tufted cells to the conditioned odor when those cells were associated with regions of focal 2-DG uptake for that odor. Mitral/tufted cells demonstrated significantly more suppressive and fewer excitatory responses to the conditioned odor than cells in control pups. Response patterns to a novel odor were not similarly modified. 4. Response patterns of mitral/tufted cells distant from the focal region of 2-DG uptake to the conditioned odor were not modified by conditioning compared with control pups. 5. The difference in response pattern between cells in the 2-DG focus and cells distant to the 2-DG focus was apparent within 500 ms of the stimulus onset. Given the respiratory rate of these pups (2 Hz), these data suggest that the modified response pattern occurred on the first inhalation of the learned odor. 6. These data demonstrate that both spatial and temporal patterns of olfactory bulb output neuron activity are used in the coding of olfactory information in the bulb. Furthermore, these spatial/temporal response patterns can be modified by early learning.

Animals↗
PubMed1999

Synchronized oscillatory discharges of mitral/tufted cells with different molecular receptive ranges in the rabbit olfactory bulb.

Individual glomeruli in the mammalian olfactory bulb represent a single or a few type(s) of odorant receptors. Signals from different types of receptors are thus sorted out into different glomeruli. How does the neuronal circuit in the olfactory bulb contribute to the combination and integration of signals received by different glomeruli? Here we examined electrophysiologically whether there were functional interactions between mitral/tufted cells associated with different glomeruli in the rabbit olfactory bulb. First, we made simultaneous recordings of extracellular single-unit spike responses of mitral/tufted cells and oscillatory local field potentials in the dorsomedial fatty acid-responsive region of the olfactory bulb in urethan-anesthetized rabbits. Using periodic artificial inhalation, the olfactory epithelium was stimulated with a homologous series of n-fatty acids or n-aliphatic aldehydes. The odor-evoked spike discharges of mitral/tufted cells tended to phase-lock to the oscillatory local field potential, suggesting that spike discharges of many cells occur synchronously during odor stimulation. We then made simultaneous recordings of spike discharges from pairs of mitral/tufted cells located 300-500 microm apart and performed a cross-correlation analysis of their spike responses to odor stimulation. In approximately 27% of cell pairs examined, two cells with distinct molecular receptive ranges showed synchronized oscillatory discharges when olfactory epithelium was stimulated with one or a mixture of odorant(s) effective in activating both. The results suggest that the neuronal circuit in the olfactory bulb causes synchronized spike discharges of specific pairs of mitral/tufted cells associated with different glomeruli and the synchronization of odor-evoked spike discharges may contribute to the temporal binding of signals derived from different types of odorant receptor.

Aldehydes↗
PubMed2002

The olfactory bulb as an independent developmental domain.

The olfactory system is a good model to study the mechanisms underlying guidance of growing axons to their appropriate targets. The formation of the olfactory bulb involves differentiation of several populations of cells and the initiation of the central projections, all under the temporal and spatial patterns of gene expression. Moreover, the nature of interactions between the olfactory epithelium, olfactory bulb and olfactory cortex at early developmental stages is currently of great interest. To explore these questions more fully, the present review aims to correlate recent data from different developmental studies, to gain insight into the mechanisms involved in the specification and development of the olfactory system. From our studies in the pax6 mutant mice (Sey(Neu)/Sey(Neu)), it was concluded that the initial establishment of the olfactory bulb central projections is able to proceed independently of the olfactory sensory axons from the olfactory epithelium. The challenge that now remains is to consider the validity of the olfactory bulb as an independent development domain. In the course of evaluating these ideas, we will review the orchestra of molecular cues involved in the formation of the projection from the OB to the olfactory cortex.

Animals↗
PubMed2002

cAMP-independent responses of olfactory neurons in Xenopus laevis tadpoles and their projection onto olfactory bulb neurons.

We report on responses of olfactory receptor neurons (ORNs) upon application of amino acids and forskolin using a novel slice preparation of the olfactory epithelium of Xenopus laevis tadpoles. Responses were measured using the patch-clamp technique. Both amino acids and forskolin proved to be potent stimuli. Interestingly, a number of ORNs that responded to amino acids did not respond to forskolin. This suggests that some amino acids activate transduction pathways other than the well-known cAMP-mediated one. The differential processing of cAMP-mediated stimuli on the one hand and amino acid stimuli on the other was further elucidated by calcium-imaging of olfactory bulb neurons using a novel nose-olfactory bulb preparation of Xenopus laevis tadpoles. The projection pattern of amino acid-sensitive ORNs to olfactory bulb neurons differed markedly from the projection pattern of forskolin-sensitive ORNs. Olfactory bulb neurons activated by amino acids were located laterally compared to those activated by forskolin, and only a small proportion responded to both stimuli. The ensemble of neurons activated by forskolin was also activated by the phosphodiesterase inhibitor 3-isobutyl-1-methylxanthine (IBMX) and the membrane-permeant cAMP analogue 8-(4-chlorophenylthio)adenosine 3',5'-cyclic monophosphate (pCPT-cAMP). We therefore conclude that sensory transduction of a number of amino acids is cAMP independent, and amino acid- and cAMP-mediated responses are processed differentially at the level of the olfactory bulb.

Amino Acids↗
PubMed1981

Volumetric comparison of the accessory olfactory bulb in bats.

Accessory olfactory bulb (AOB) volumes were calculated in 26 chiropteran species. The percentage of the AOB in relation to the olfactory bulb complex ranges between 0.18 and 2.70%. The allometric comparison reveals the high development of this structure in the phyllostomatid subfamily of Glossophaginae. Carolliinae and Sturnirinae are near the chiropteran average. Smaller AOBs can be observed in Sphaeronycteris, Diphylla, Pteronotus parnellii and Miniopterus schreibersi. Comparison of the right/left AOB size in 56 specimens showed a slight predominance of the latter (average 50.6% of total volume). Intraspecific variability of this structure is pronounced, reaching a maximum SEM of 23.2% in Artibeus lituratus. Other species with well-developed AOBs tend to show high intraspecific variability. The role of the accessory olfactory system is seen by recent investigators mainly in its participation in sexual behavior and in close connection with the 'Flehmen' gesture, which is predominantly shown by male individuals. When comparing the size of the AOB in 18 bat species where both sexes were represented, only half of them showed smaller AOBs in females. This demonstrates that there is no tendency towards reduction of this structure in females.

Animals↗
PubMed2003

Microglia in the olfactory bulb of rats during postnatal development and olfactory nerve injury with zinc sulfate: a lectin labeling and ultrastrucutural study.

Using isolectin (GSA I-B4) as a marker, this study examined the possible alterations of lectin-labeled membranous glycoproteins in microglial cells in the olfactory bulb of normal development and under experimentally induced degeneration. In light microscopy, several morphological types of microglial cells representing different degrees of cell differentiation were distributed in the bulb laminae. A gradient of microglial differentiation extending from the intermediate to superficial and intermediate to deep occurs in the bulb layers. The differentiation gradient and lectin labeling pattern of microglial cells in the developing bulb resembled those in other areas of the brain tissues. Differentiating microglia showed a gradual diminution of lectin staining when the nascent round cells transformed into the mature ramified cells. Microglia in the external plexiform layer of the olfactory bulb were the first to mature and the cells expressed very weak lectin reactivity. In mature or adult rats, some microglial cells showing intense lectin labeling were observed in the olfactory nerve layer, granule cell layer and subependymal layer. Ultrastructurally, lectin labeling was localized at the trans saccules of the Golgi apparatus. Microglial cells in other bulb laminae, however, exhibited a negative reaction for the isolectin at the Golgi apparatus. Following intranasal irrigation of zinc sulfate, some microglial cells in the olfactory nerve layer and glomerular layer were activated to become phagocytic cells with increased lectin labeling at their ramified processes. GSA I-B4 staining was also localized at their trans saccules of the Golgi apparatus. The lectin labeling pattern of these phagocytic cells resembled that of differentiating microglia in postnatal bulbs, suggesting that bulb microglia in the lesioned sites were activated through cell dedifferentiation into macrophages.

Animals↗
PubMed1995

Neurotransmitter release in the accessory olfactory bulb during and after the formation of an olfactory memory in mice.

Female mice form an olfactory memory to the pheromones of the mating male during a critical period after mating. Previous experiments have shown that the neural changes underlying this memory are located in the accessory olfactory bulb, are dependent on noradrenergic neurotransmission, and most likely involve changes at the mitral-granule cell reciprocal synapses. Using the technique of in vivo microdialysis we have followed changes in a range of neurotransmitters before, during and after memory formation. The increase in GABA levels in response to a glutamate challenge was greater during and after memory formation than before. The aspartate/GABA ratio was decreased following memory formation, during exposure to the pheromones of the mating male. These findings are consistent with our hypothesis that memory formation involves a long-lasting increase in the inhibition of the subset of mitral cells that respond to the mating male's pheromones. Unexpectedly, there were increases in the concentrations of the excitatory transmitters glutamate and aspartate in non-mating females, immediately following male exposure, and two days later in response to re-exposure to the same male pheromones. These results suggest that exposure to male pheromones alone, without the association of mating, causes a long-lasting decrease in the inhibitory control of the subset of mitral cells responding to these pheromones. The implication of these results is that two types of synaptic plasticity can occur in the accessory olfactory bulb. The association of mating and pheromonal exposure induces memory formation by increasing the inhibition of the pheromonal signal at the level of the accessory olfactory bulb, thereby preventing them from activating the neuroendocrine block to pregnancy. Male exposure without mating appears to have the opposite effect, decreasing the inhibition of the pheromonal signal and potentiating the oestrous-inducing effects of the male pheromones.

Animals↗
PubMed1981

Aging in the rat olfactory system: correlation of changes in the olfactory epithelium and olfactory bulb.

Two regions closely linked synaptically (olfactory epithelium and olfactory bulb) have been compared in an age-graded series of rats. Previous findings of growth and atrophy of constituent elements in Sprague-Dawley Wisconsin (SD) rats have now been confirmed in Charles River (Crl) rats. In the olfactory bulbs of Charles River (Crl) rats, the volume of layers, the number of olfactory axodendritic synapses in the glomeruli, the total volume of glomerular dendrites, and the size of mitral cell bodies all approximately double between 3 and 24--27 months, and then all decrease by 36 months. Unlike SD rats, however, no loss in the number of mitral cells occurs in Crl rats, and the increase in volume of the olfactory bulbs from 3 to 24 months is approximately double that of SD rats. In the olfactory epithelium the total number of septal olfactory receptors more than doubles between 3 and 18--24 months and then declines markedly, as does the volume of olfactory axons in olfactory bulb glomeruli. Comparison of the regression lines for change in number of septal receptors with that of the size of mitral cell bodies discloses that the decline in number of receptors begins several months earlier than the decline in mitral cell size. This suggests that the atrophic changes in the olfactory bulb may in part be secondary to changes in the receptors of the olfactory epithelium. Numbers of synapses in the glomeruli appear to decline less markedly with age than the number of receptors, and a significant increase in number of synapses per receptor occurs in the oldest group studied (33 months), suggesting a compensatory increase in the relative number of synapses per receptor in the surviving receptors.

Aging↗
PubMed2003

Activation of the cyclic AMP response element-binding protein signaling pathway in the olfactory bulb is required for the acquisition of olfactory aversive learning in young rats.

Long-term memory formation requires both gene expression and protein synthesis. Phosphorylation of the transcription factor cyclic AMP response element-binding protein (CREB) is thought to be important in processes underlying long-term memory. To clarify the role of CREB in olfactory aversive learning in young rats, we carried out behavioral pharmacology and Western blot analyses. On postnatal day 11, oligodeoxynucleotides were infused directly into the bilateral olfactory bulbs through cannulae implanted prior to training in a classical conditioning paradigm with citral odor and foot shock. On the following day the odor preference test was performed. After training, saline-infused animals spent significantly shorter time over the citral odor zone. Infusion of CREB antisense oligodeoxynucleotides 6 h before or during training, however, prevented olfactory aversive learning without affecting memory retention 1 h after training. CREB scrambled oligodeoxynucleotides infusions had no effect on olfactory learning. When infused 6 h after training, none of oligodeoxynucleotides had an effect on time spent over the odor zone. Using Western blotting, we analyzed CREB in nuclear extracts obtained from the young rats after training. Marked increases in phosphorylated CREB were sustained from 10 to 360 min after the odor-shock pairing in animals which were subjected to both, in comparison with levels 30 min in animals which were subjected to odor only or no stimulation. Total CREB levels showed no differences among groups. Infusion of CREB antisense oligodeoxynucleotides significantly reduced the expression of phosphorylated and total CREBs in the olfactory bulb. These results show that the synthesis and phosphorylation of CREB are required for the acquisition of olfactory aversive learning in young rats, and that this requirement for the CREB signaling pathway has a critical time window.

Animals↗
PubMed1997

Carnosine-induced inward currents in rat olfactory bulb neurons in cultured slices.

Olfactory neurons are rich in carnosine, which is regarded as a possible neurotransmitter between olfactory neurons and the olfactory bulb neurons. In the present study, we used the organotypic slice culture technique to culture olfactory bulb neurons and explore carnosine's role in the olfactory system. Inward current responses to carnosine upon increases in membrane conductance were recorded under whole-cell voltage-clamp conditions. The inward currents were desensitized during continuous application of carnosine. At carnosine concentrations above 50 microM, the current responses increased with increases in carnosine concentrations. The present results suggest that carnosine serves as an excitable neuroeffector between olfactory neurons and olfactory bulb neurons.

Animals↗
PubMed1992

Emergent properties of odor information coding in a representational model of the salamander olfactory bulb.

In the salamander olfactory bulb, mitral output cells exhibit a variety of responses to electrical and odor stimulation, but the cellular interactions within the bulb that give rise to these responses are not completely understood. We have developed a computer model to investigate whether available data are sufficient for formulating a simulated bulb circuit that can generate realistic mitral cell output. A set of coupled difference equations incorporating mathematical descriptions of anatomical and physiological data was used to calculate changes in membrane potentials of olfactory bulb neurons over time. Model mitral cells showed responses to simple orthodromic and antidromic electrical stimuli that were similar to salamander intracellular responses. Without changing the parameters of the equations, simulated odor stimuli were applied that elicited complex patterns of mitral depolarization, spike activation, and hyperpolarization that emerged from the interactions among the numerous elements in the model. As with the electrical stimuli, model mitral responses to odor were also strikingly similar to those of real mitral cells. As an initial test of how different circuit components contribute to the responses, the lateral interactions between mitral cells and bulbar interneurons were manipulated. Tests with reduced lateral interactions and other tests with no inhibitory synaptic connections both produced mitral cell outputs that were uncharacteristic of salamander recordings. The similarity of the model's output to the complex properties of salamander single-cell recordings suggests that several critical features of the bulb circuit responsible for shaping mitral cell responsivity have been captured.(ABSTRACT TRUNCATED AT 250 WORDS)

Ambystoma↗
PubMed2002

Release-independent short-term facilitation at GABAergic synapses in the olfactory bulb.

Neurones of the olfactory bulb are innervated by GABA-releasing axons and dendrites of diverse origin. Here, I studied GABAergic neurotransmission in juxtaglomerular cells using whole-cell voltage-clamp recordings in acute olfactory bulb slices. Spontaneous IPSCs were fully blocked by the GABA(A) receptor antagonist SR95531 (40 microM) and the sodium channel blocker tetrodotoxin (1 microM). The IPSCs had mean amplitudes of 125+/-86 pA and relatively slow biexponential decay times (tau(1)=4.3+/-1.0 ms (67+/-12%), tau(2)=16.9+/-2.7 ms) at physiological temperatures. Short-term plasticity of evoked IPSCs showed two distinct patterns: depressing (n=4 cells) and facilitating-depressing (n=9). In two cells, postsynaptic responses were mediated by single functional release sites. During a train of stimuli (4 stimuli at 20 Hz), the release probability increased by two-fold, whereas the potency (postsynaptic responses excluding failures) decreased by ~15%. The increase in release probability for the second stimulus in the train also occurred when the first action potential failed to release transmitter. However, the decrease in the potency was only observed if the preceding action potential released transmitter. These results reveal a heterogeneity in the short-term plasticity of evoked IPSCs in juxtaglomerular cells and demonstrate that the short-term facilitation at some GABAergic synapses is independent of release.

Algorithms↗