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Taurine action on mitral cell activity in the frog olfactory bulb in vivo.

Taurine (TAU) is a free amino acid that is particularly abundant in the olfactory bulb. In the frog, TAU is located in the terminations of the primary olfactory axons and in the granular cell layer. TAU action seems to be associated with gamma amino butyric acid (GABA), the main inhibitory neurotransmitter involved in the processing of the sensory signal. The present study was designed to assess the action of TAU in vivo during the olfactory network's stimulation by odors. It was performed by recording the single-unit activity of mitral cells, the main bulbar output neurons. TAU effects were tested on both their spontaneous and odor-induced firing activity. Interactions between TAU and GABA were examined by analyzing TAU effects under the selective blocking action of GABAA or GABAB antagonists. TAU was found to suppress the spontaneous firing of mitral cells, mainly without altering their odor response properties. By testing GABA antagonists, we further show that TAU action is associated with GABAergic inhibitory mechanisms mainly via GABAB receptors. Thus, TAU action clearly reduces background activity in favor of the emergence of the odor-induced activity in the same manner as GABA action does via GABAB receptors. As a conclusion, we propose that, in the frog olfactory bulb, the joint actions of TAU and GABA may favor the processing of the primary sensory information by increasing the signal to noise ratio.

Action Potentials↗

The influence of training on chemosensory event-related potentials and interactions between the olfactory and trigeminal systems.

It is not possible to accurately predict the perceptual response to odorants and odorant mixtures without understanding patterns of suppression and facilitation that result from interactions between the olfactory and trigeminal systems. The current study extends previous findings by exploring the effect of intensive training on the interaction between these systems and also by using a different mixed chemosensory stimulus to examine whether the principles established in earlier studies generalize to different odorants. Stimuli were chosen so as to selectively activate the olfactory (H2S) and trigeminal (CO2) nerves. In addition, linalool was included as a stimulus that activated both systems. Thirty-five participants (19 men, 16 women) rated the intensity of each stimulus when presented both alone and in binary mixtures (linalool + H2S, and linalool + CO2). Chemosensory event-related potentials were obtained from three recording positions. Analysis of intensity ratings showed that linalool was significantly less intense than the other stimuli when presented alone. In binary mixtures, H2S was strongly suppressed by linalool. One week of intensive odor training produced significant and specific reductions in the intensity of linalool and H2S, both alone and in their mixture. Training with a different odor (champignol) had no effect. Chemosensory event-related potential data confirmed previous findings showing changes in topographical distribution that reflected the degree of trigeminal activity. Binary mixtures generally produced larger amplitudes than single stimuli. Latencies clearly differentiated between the three single stimuli and the binary mixtures. Changes were observed in event-related potentials that reflected those obtained for intensity ratings in that they were observed for linalool and H2S in the linalool trained group only. The amplitude of the late 'endogenous' component (P3) was significantly decreased for these odors at frontal recording sites. In summary, strong and specific training effects were observed in intensity ratings for participants trained with the test odor (linalool), but not for those trained with a different odor. This was supported by a significant decrease of amplitudes of the event-related potentials at frontal recording sites following training with the test odor only

Acyclic Monoterpenes↗

OCAM reveals segregated mitral/tufted cell pathways in developing accessory olfactory bulb.

Two functional subsets of vomeronasal sensory neurons project their axons to two segregated zones in the accessory olfactory bulb (AOB). Using immunohistochemical methods with antibodies against the novel cell adhesion molecule OCAM, we provide evidence that the segregation of functional pathways is maintained at the level of mitral/tufted (M/T) cells of the mouse AOB and that this pattern emerges early in ontogeny. During embryonic and postnatal development OCAM was strongly expressed by M/T cells in the caudal zone of the AOB where OCAM-negative vomeronasal axons terminated. In contrast, rostral zone M/T cells innervated by OCAM-positive vomeronasal axons displayed no or faint OCAM immunoreactivity. Differential expression of OCAM in segregated M/T cell pathways suggests that OCAM may be involved in defining compartments of connectivity and setting up functional subdivisions in the developing AOB.

Aging↗

Zonal organization of the mammalian main and accessory olfactory systems.

Zonal organization is one of the characteristic features observed in both main and accessory olfactory systems. In the main olfactory system, most of the odorant receptors are classified into four groups according to their zonal expression patterns in the olfactory epithelium. Each group of odorant receptors is expressed by sensory neurons distributed within one of four circumscribed zones. Olfactory sensory neurons in a given zone of the epithelium project their axons to the glomeruli in a corresponding zone of the main olfactory bulb. Glomeruli in the same zone tend to represent similar odorant receptors having similar tuning specificity to odorants. Vomeronasal receptors (or pheromone receptors) are classified into two groups in the accessory olfactory system. Each group of receptors is expressed by vomeronasal sensory neurons in either the apical or basal zone of the vomeronasal epithelium. Sensory neurons in the apical zone project their axons to the rostral zone of the accessory olfactory bulb and form synaptic connections with mitral tufted cells belonging to the rostral zone. Signals originated from basal zone sensory neurons are sent to mitral tufted cells in the caudal zone of the accessory olfactory bulb. We discuss functional implications of the zonal organization in both main and accessory olfactory systems.

Animals↗

Acquiring symptoms in response to odors: a learning perspective on multiple chemical sensitivity.

In this chapter, a learning account is discussed as a potential explanation for the symptoms in multiple chemical sensitivity. Clinical evidence is scarce and anecdotal. A laboratory model provides more convincing results. After a few breathing trials containing CO2-enriched air as an unconditioned stimulus in a compound with harmless odor substances as conditioned stimuli, subjective symptoms are elicited and respiratory behavior is altered by the odors only. Also, mental images can become conditioned stimuli to trigger subjective symptoms. The learning effects cannot be explained by a response bias or by conditioned arousal, and they appear to involve basic associative processes that do not overlap with aware cognition of the relationship between the odors and the CO2 inhalation. Learned symptoms generalize to new odors and they can be eliminated in a Pavlovian extinction procedure. In accordance with clinical findings, neurotic subjects and psychiatric cases are more vulnerable to learning subjective symptoms in response to odors. Consistent with a learning account, cognitive-behavioral treatment techniques appear to produce beneficial results in clinical cases. Several criticisms and unresolved questions regarding the potential role of learning mechanisms are discussed.

Administration, Inhalation↗

Reafference and attractors in the olfactory system during odor recognition.

Olfactory bulb activity has been postulated to be chaotic, as measured in the EEG, and to be subject to an attractor with many "wings" enabling classification of different learned odor classes. Two parallel questions are thus addressed by the work presented here: (1) what is the evidence for attractors in the olfactory system, which can mediate learned odor classes? and (2) how does the olfactory system enter a specific attractor or attractor wing associated with the learned odor during the classification process? Both of these questions address the wider notion of endogenous activity preparing the system for an expected stimulus, which is at the basis of the reafference principle. By viewing the brain as a distributed complex dynamical system with global attractors, these questions can be answered together. Rats were implanted with bipolar macroelectrodes in the Olfactory Bulb (OB), Prepyriform Cortex (PPC), Entorhinal Cortex (EC), and Dentate Gyrus (DG), and then trained in an operant paradigm to press a bar for a reward in the presence of one odor and to receive no reward in the presence of another odor. Local Field Potentials (LFP) were recorded simultaneously from the structures during the operant task. We present evidence for three endogenous events: (1) preafference, which is manifested both by the EC entering an attractor and a mid-range signal (15-30 Hz) which appears to be passed from the EC to the OB just before the OB enters an attractor; (2) afference, where the OB enters an attractor during the odor recognition period of the experiment and the LFP recordings indicate that the OB drives the other structures in all frequency bands, especially the high gamma band (65-100 Hz) associated with the OB burst frequency; and (3) reafference or post-afference, which is accompanied by a lower frequency gamma band signal (40-60 Hz) originating in the PPC and passed to both the OB and the EC just before the onset of the motor response to the odor. We use a new method, NECTAR (Nonparametric Exact Contingency Table Association Routine), related to mutual information, to verify what is seen with coherence and phase estimates, the apparent driving of each structure at different times in the odor trials, and to display evidence for non-periodic attractors governing both individual physiological structures and the system of structures. This is the first evidence of an endogenous, limbic event associated with sensory perceptual tuning in a mammal. These results are also the first experimental confirmation that the attractors governing olfactory activity involve multiple sites in the olfactory/limbic system and implement the process of attention.

Adaptation, Physiological↗

Spatiotemporal analysis of prepyriform, visual, auditory, and somesthetic surface EEGs in trained rabbits.

1. Spatial ensemble averages were computed for 64 traces of electroencephalograms (EEGs) simultaneously recorded from 8 x 8 arrays over the epidural surfaces of the prepyriform cortex (PPC) and visual, somatic, and auditory cortices. They revealed a common waveform across each array. Examination of the spatial amplitude modulation (AM) of the waveform revealed classifiable spatial pattern in short time segments. The AM patterns varied within trials after presentation of identical conditioned stimuli, and also between trials with differing stimuli. 2. PPC EEGs revealed strong correlates with the respiratory rhythm; neocortical EEGs did not. 3. Time ensemble averaging of the PPC EEG attenuated the oscillatory bursts, indicating that olfactory gamma oscillations (20-80 Hz) were not phase-locked to the times of stimulus delivery but instead to inhalations. Time ensemble averages of neocortical recordings across trials revealed average evoked potentials starting 30-50 ms after the arrival of the stimulus. 4. Average temporal fast Fourier transform (FFT) power spectral densities (PSDs) from pre- and poststimulus PPC EEG segments revealed a peak of gamma activity in olfactory bursts. 5. The logarithm of the average temporal FFT PSDs from pre- and poststimulus neocortical EEG segments, when plotted against log frequency, revealed 1/f-type spectra in both pre- and poststimulus segments for negative/aversive conditioned stimuli (CS-) and positive/rewarding conditioned stimuli (CS+). The alpha'- and beta'-coefficients from the regression of Eq. 2 onto the average PSDs were significantly different between pre- and poststimulus segments, owing to the evoked potentials, but not between CS- and CS+ stimulus segments. 6. Spatiotemporal patterns were invariant over all frequency bins in the 1/f domain (20-100 Hz). Spatiotemporal patterns in the 2- to 20-Hz domain progressively differed from the invariant patterns with decreasing frequency. 7. In the spatial frequency domain, the logarithm of the average spatial FFT power spectra from pre- and poststimulus neocortical EEG segments, when plotted against the log spatial frequency, fell monotonically from the maximum at the lowest spatial frequency, downwardly curving to a linear 1/f spectral domain. This curve in the 1/f spectral domain extended from 0.133 to 0.880 cycles/mm in the PPC and from 0.095 to 0.624 cycles/mm in the neocortices. 8. Methods of FFT and principal component analysis (PCA) EEG decomposition were used to extract the broad-spectrum waveform common to all 64 EEGs from an array. AM patterns for the FFT and PCA components were derived by regression. They were shown by cross-correlation to yield spatial patterns that were equivalent to each other and to AM patterns from calculation of the 64 root-mean-square amplitudes of the segments. 9. Each spatial AM pattern was expressed by a 1 x 64 column vector and a point in 64-space. Similar patterns formed clusters, and dissimilar patterns gave multiple clusters. A statistical test was devised to evaluate dissimilarity by a Euclidean distance metric in 64-space. 10. Significant spatial pattern classification of CS- versus CS+ trials (below the 1% confidence limit for 20 of each) was found in discrete temporal segments of poststimulus data after digital temporal and spatial filter optimization. 11. Varying the analysis window duration from 10 to 500 ms yielded a window length of 120 ms as optimal for pattern classification. A 120-ms window was subsequently stepped across each record in overlapping intervals of 20 ms. Windows in which episodic, significant CS+/CS- differences occurred lasted 50-200 ms and were separated by 100-200 ms in the poststimulus period. 12. Neocortical spatial patterns changed under reinforcement contingency reversal, showing a lack of invariance in respect to stimuli and a dependence on context and learning, as previously found for the olfactory bulb and PPC.

Animals↗

Perceptual and neural olfactory similarity in honeybees.

The question of whether or not neural activity patterns recorded in the olfactory centres of the brain correspond to olfactory perceptual measures remains unanswered. To address this question, we studied olfaction in honeybees Apis mellifera using the olfactory conditioning of the proboscis extension response. We conditioned bees to odours and tested generalisation responses to different odours. Sixteen odours were used, which varied both in their functional group (primary and secondary alcohols, aldehydes and ketones) and in their carbon-chain length (from six to nine carbons). The results obtained by presentation of a total of 16 x 16 odour pairs show that (i) all odorants presented could be learned, although acquisition was lower for short-chain ketones; (ii) generalisation varied depending both on the functional group and the carbon-chain length of odours trained; higher generalisation was found between long-chain than between short-chain molecules and between groups such as primary and secondary alcohols; (iii) for some odour pairs, cross-generalisation between odorants was asymmetric; (iv) a putative olfactory space could be defined for the honeybee with functional group and carbon-chain length as inner dimensions; (v) perceptual distances in such a space correlate well with physiological distances determined from optophysiological recordings of antennal lobe activity. We conclude that functional group and carbon-chain length are inner dimensions of the honeybee olfactory space and that neural activity in the antennal lobe reflects the perceptual quality of odours.

Afferent Pathways↗

[Characteristics of the reaction of submissive mice to distant zoosocial stimuli].

In aggressive and submissive mice well-experienced in victories and defeats, the reaction was studied to another male in neighbour section of "home" cage divided by transparent partition with holes. The activity of submissive mice near the partition was significantly lower than that of control animals without experience of agonist contacts or aggressive. Submissive mice did not react to the appearance of unknown smell in the neighbouring section while the animals of other studied groups sharply increased their activity near the partition. It is suggested that long experience of defeat leads to inhibition of perception of specific zoosocial (olfactory) stimuli triggering manifestation of aggressive reaction to another individual.

Aggression↗

[Multipolar electrical stimulation of the olfactory bulb as a model for studying spatial basis of olfactory coding].

Low-intensity electrical stimulation of the olfactory bulb through a set of spatially distributed electrodes was used as a model in order to study the spatial basis of the olfactory coding, in rats. The results indicate that this multipolar stimulation can serve as a conditional stimulus in a learned aversion paradigm; in addition, water-deprived animals displayed high ability to discriminate and memorize various combinations of electrode positions paired with differently palatable solutions.

Animals↗

[Comparative volumetric analysis of the principal subdivisions of the telencephalon in saurian reptiles].

The volumetric measure of the main subdivisions of the telencephalon has been carried on 24 species of Lizards and 2 species of Snakes. The studied structures are termed as follows: main and accessory olfactory bulbs, medial cortex (M 1 and M 2), dorsal cortex (D 1, D 2 and D 3), lateral cortex (L), Septum, Tuberculum olfactorium, dorsal and ventral striatum, amygdala and nucleus sphaericus. The analysis of the datas makes use of the SNEL L's formula which relates the volume of the various telencephalic subdivisions (V) to the somatic weight (S): V = k x S alpha. Each alpha value is compared to the value of the coefficient of allometry (A) of the whole brain. The evolutive (phylogenetic) growth of a structure is said fast (or slow) when its corresponding alpha value is higher (or lower) than the encephalic A value. At the cortical level such analysis shows the progressive nature of the dorsal cortex. A partition of the sample into Lacertomorpha (14 species) and Dracomorpha (10 species) (in agreement with the NORTHCUTT'S definition of his Type I and Type II Lizards) corroborates this cortical detail, more distinctly with the second group as well (especially for the D 2 portion). Moreover the high number of progressive structures among the Dracomorpha leads to consider this group as phylogenetically the most advanced in the Order of Lizards. The somatic indices are calculated according the allometric characteristics of the Reference Lizards. The judicious choice of some species allows to show how the development of a biological function may be expressed by the values of the indices of the related structures. For examples: dorsal cortex, dorsal striatum and mode of locomotion; olfactory bulbs, lateral cortex, part M 1 of the medial cortex and olfactory system; D 3 subdivision of the dorsal cortex and visual performances. The duality between Lacertomorpha and Dracomorpha is therefore corroborated by significant differences found for the various indices of a great number of telencephalic subdivisions. It leads moreover to find, grosso modo, two functional types of Lizards: moving-on-the-ground and wellsmelling (mainly Lacertomorpha) on the one hand, arboreal and with a fine vision (mainly Dracomorpha) on the other hand. The isoponderal percentages take an useful illustration of these results; it allows to establish the telencephalic pattern of a standard Lizard in which the pallium keeps the larger part (42%); in the pallium itself, the M 1 subdivision of the medial cortex has the most important percentage, a little more than the D 2 part of the dorsal cortex. A comparative study carried on 2 Snakes gives for Boa constrictor the lowest values of the indices, for almost all the structures. In return Natrix natrix stays, for a great number of structures, close to the level of the legless Lizards; this last result confirms distinctly the two levels of telencephalization already found in Snakes (PLATEL, 1976 a).

Amygdala↗

[Smell perception in smokers].

The author studied the changes of olfactory function in smokers. He established the fact that the threshold of the perception of smell increases in smokers. By studying the subjective experience of the intensity of smell he established an appearance which he named olfactory recruitment, the increased growth of smelling sensitivity, which points out the changes, not only on the olfactive mucous membranes, but on the olfactive paths as well. The author concludes that smoking influences the decrease of the function of smell and emphasizes the necessity of advising the stopping of smoking in the treatment of all types of smelling disorders, as one of the important elements of treatment.

Humans↗

Odor perception and beliefs about risk.

Although the perceptual response to environmental odors can be quite variable, such variation has often been attributed to differences in individual sensitivity. An information-processing analysis of odor perception, however, treats both the reception and the subsequent evaluation of odor information as determinants of the perceptual response. Two experiments investigated whether a factor that influenced the evaluation stage affected the judgement of odor quality and the degree of adaptation to the odor. People were surveyed in order to measure their tacit perceptions of the healthfulness or hazardousness of nine common olfactory stimuli, and the instructional context influenced quality perception. In a second experiment subjects were exposed to an ambient odor under one of three different conditions, and odorant characterization influenced the degree of adaptation to the odor. Subjects who were led to believe the odor was a natural, healthy extract showed adaptation; those told that the odor was potentially hazardous showed apparent sensitization; while those told that the odor was a common olfactory test odorant showed a mixed pattern: some exhibited adaptation, whereas others showed sensitization. However, detection thresholds obtained before and after exposure showed adaptation effects that are characteristic of continuous exposure. These findings raise the possibility that cognitive factors may be modulating the overall sensory perception of odor exposure (i) for some individual who exhibit extreme sensitivity to odors and (ii) in situations where adaptation to environmental odors is expected but does not occur.

Adolescent↗