Repeated subinflammatory ultraviolet B irradiation increases substance P and calcitonin gene-related peptide content and augments mustard oil-induced neurogenic inflammation in the skin of rats

The objective of the present study was to investigate interactions between odor hedonic tone, perceived odor intensity, olfactory judgments and stimulated nostril laterality. Subjects were stimulated in the right and the left nostril separately, with two odors: vanillin (pleasant); and indole (unpleasant). They had two tasks to perform: an intensity judgment; and an affective judgment. Two concentrations (one strong and one weak) of each odor were presented. Odors were presented for a short period corresponding to one inhalation (about 1 s). The inter-stimulus interval was always 30 s. The nostril stimulated and task presentation order were counterbalanced according to a Latin square. Odor presentation order was randomized for each subject. Results indicated that response times for unpleasant odors were significantly shorter than for pleasant odors during affective judgment and right nostril stimulation, indicating greater right hemisphere efficiency for the decoding of unpleasant affects induced by odors.     

Emotional reactivity to odors: Olfactory sensitivity and the span of emotional evaluation separate the genders

Numerous studies report gender differences in emotional reactivity in health and disease and the perception of odors is closely linked to the limbic system. In order to investigate gender differences in the emotional perception of odors we extended the Sniffin’ Stick Test with analogue rating scales for hedonic (pleasantness/unpleasantness) and intensity estimates. We matched 172 healthy subjects (86 females and 86 males) on age in order to balance the study population for three age strata (A: 19–39 years, B: 40–59, C: 60 years and above). Overall odors in our statistical analysis demonstrated significant gender differences for the absolute hedonic estimates but not for the relative hedonic and not for the intensity estimates. These findings demonstrate that women evaluate the pleasantness of perceived odors in a more extreme manner than men without significant differences in hedonic polarity (pleasantness/unpleasantness). Thus, we report a singular significant effect of gender on the dimension valence (hedonic estimation) independent from the dimension intensity. Our findings are in accordance with gender differences in facial reactivity to auditory stimuli and differences in the evaluation of emotional pictures. Investigating olfactory sensitivity females detected n-butanol and discriminated the 16 odors of the test significantly better than males. These results indicate that females possess higher olfactory sensitivity.     

The role of temporal lobe and orbitofrontal cortices in olfactory memory function.

Differences in verbal and nonverbal olfactory identification and recognition were examined among three groups with brain impairment. A left cerebrovascular accident (LCVA) group, a right CVA (RCVA) group, and a traumatic brain injury (TBI) group were compared with two nonimpaired age-matched comparison groups on olfaction identification and recognition abilities. Odors were presented to the left and right nostrils, which maximized hemispheric differences in olfactory processing. Results showed that persons with LCVA demonstrated the greatest impairment on the verbal identification of odors, while persons with RCVA showed the most impairment on the nonverbal identification of odors. Persons with TBI showed an inconsistent impairment across both verbal and nonverbal odor identification tasks. Odor recognition was impaired in both CVA groups as well. In contrast, persons with TBI performed better on the delayed odor recognition tasks. Results are discussed in relation to hemispheric differences in processing olfactory information.     

Differences in naming accuracy of odors presented to the left and right nostrils

Two interlinked areas of debate within psychology are the existence of hemispheric specialization for olfactory processing and whether odors are processed primarily as perceptual codes or as a set of semantic features. This study compared accuracy in naming and judgments of familiarity by right handed subjects to common odors that were presented to the left or right nostrils. There was significantly better production of the correct name of the odor after left side presentation, than after right. Familiarity ratings were identical.     

Olfactory abnormalities in anxiety disorders

The olfactory system plays an important role in both animal and human anxiety reactions. However, results on olfactory performance during the course of clinical anxiety disorders remain scarce. Therefore, we conducted an exploratory pilot study in 17 patients currently diagnosed with an anxiety disorder. Patients participated in olfactory and psychological testing and were compared to 17 healthy controls. Statistical analyses revealed significant deficits concerning olfactory discrimination in patients, while no changes in threshold and identification ability occurred. Most interestingly, patients showed higher intensity estimates and an increased rating range concerning olfactory hedonic judgements. Results are discussed in light of prior findings and related to neural correlates of olfactory perception and anxiety reactions.     

Functional anatomy of human odor sensation,discrimination, and identification in health and aging

Aging of cerebral olfactory regions was studied in 5 younger and 6 older healthy adults, matched by odor discrimination and identification scores, with positron emission tomography during odor sensory stimulation, discrimination, and identification tasks. Sensory stimulation engaged bilateral piriform and orbitofrontal regions, but neither discrimination nor identification evoked added temporal or orbital activity. Discrimination involved the hippocampus, implicating its role in serial odor comparisons (olfactory working memory). Left inferior frontal activity during identification may reflect semantic associations. Older participants deactivated the left gyrus rectus/medial orbital gyrus (GR/MOG) during sensory stimulation but activated GR/MOG during discrimination and identification. Adjusting for detection threshold eliminated GR/MOG group differences during sensory stimulation. Diminished threshold may lead to reduced engagement of olfactory association areas.     

Cross-modal integration between odors and abstract symbols

This study aimed to investigate the cross-modal association of an “abstract symbol,” designed for representation of an odor, with its corresponding odor. First, to explore the associations of abstract symbols with odors, participants were asked to match 8 odors with 19 different abstract symbols (Experiment 1). Next, we determined whether congruent symbols could modulate olfactory perception and olfactory event-related potentials (ERPs) (Experiment 2). One of two odors (phenylethanol (PEA) or 1-butanol) was presented with one of three conditions (congruent or incongruent symbol, no-symbol), and participants were asked to rate odor intensity and pleasantness during olfactory ERP recordings. Experiment 1 demonstrated that certain abstract symbols could be paired with specific odors. In Experiment 2 congruent symbol enhanced the intensity of PEA compared to no-symbol presentation. In addition, the respective congruent symbol increased the pleasantness of PEA and the unpleasantness of 1-butanol. Finally, compared to the incongruent symbol, the congruent symbol produced significantly higher amplitudes and shorter latencies in the N1 peak of olfactory ERPs. In conclusion, our findings demonstrated that abstract symbols may be associated with specific odors.     

Identification of single dissimilar odors is achieved by humans with a single sniff

The duration that a single odor needs to be sniffed for identification was determined for 18 humans. A hot wire anemometer and an oscilloscope were used to monitor the duration, volume and inhalation rate of sniffs. In Experiment 1 subjects used 1, 3 or 5 natural sniffs, or an unlimited number of natural sniffs to sample seven dissimilar single odors of moderate perceived intensity, and demonstrated that each odor could be identified with a single sniff. In Experiment 2 subjects demonstrated that each of the odors could be identified with the shortest sniff (0.42 sec) they could physically achieve. In Experiment 3 tests with two of the odorants at several concentrations showed that sniff duration influences identification over a narrow range of concentrations that is just above the recognition threshold. These results together with earlier data that described the optimum conditions for the detection of an odor and the perception of odor intensity, provide information that is necessary for the development of a standard olfactometer and standard methods for human olfactory measurements.     

Working memory across nostrils

Whether olfactory working memory involves verbal representations or neural images of odor per se remains unclear. This study investigated whether verbal representation influences performance in an olfactory delayed-match-to-sample task and used monorhinal presentation to generate hypotheses as to the underlying anatomy of this mechanism. The main findings were that (a) nameable odorants were easier to remember than hard-to-name odorants and (b) the nameability effect was more pronounced when the evaluation was done across nostrils. Considering these results within a proposed model implies dual representation in olfactory working memory: All odors, nameable and hard to name, are represented both perceptually and verbally.     

Odor-reward learning and enrichment have similar effects on odor perception

We compared the respective effects of odor enrichment and reward-driven discrimination learning on the perception of odors. Experimental rats were exposed to two odorants for 1-h periods once daily over 10 days, or trained with the same two odorants in a forced-choice discrimination task during a daily 20 trial session over 10 days. Spontaneous discriminations between pairs of chemically similar odors were tested before and after the odor enrichment period using an olfactory habituation/discrimination task. We found that both enrichment and discrimination learning improved the subsequent perception of other, chemically unrelated odorants in similar manners. These results show that odor experience, whether passive or active, changes perception in the manner predicted based on other groups' electrophysiological experiments.     

Neuroanatomical correlates of olfactory performance

We investigated associations between olfactory function and gray matter thickness in 46 healthy young subjects by means of an automated technique for measuring cortical thickness. We used an extended version of the Sniffin’ Sticks test to assess olfactory function, including odor threshold, concentration discrimination, quality discrimination, and odor identification. We observed a correlation between olfactory performance and cortical thickness of structures involved in earlier and later stages of chemosensory processing such as right medial orbitofrontal cortex, right insula, and adjacent cortex. Furthermore, we found significant bilateral correlations of olfactory performance with cortical thickness of areas around the central sulcus bilaterally, structures responsible for voluntary respiration and sniffing. In addition to expected general sex effects on cortical thickness, we observed areas, such as the entorhinal cortex, occipital cortex, intraparietal sulcus and insula (all in the right hemisphere), where the correlation between higher order olfactory functions and cortical thickness differed between women and men. These data demonstrate, for some neuroanatomical structures, a link between cortical thickness and olfactory function, in that thicker cortex is usually associated with better performance, but not always. This association between anatomy and olfactory performance suggests a possible biological explanation for the high degree of individual differences and sex effects observed in higher order olfactory tasks.     

Functional mapping of human brain in olfactory processing: a PET study

This study describes the functional anatomy of olfactory and visual naming and matching in humans, using positron emission tomography (PET). One baseline control task without olfactory or visual stimulation, one control task with simple olfactory and visual stimulation without cognition, one set of olfactory and visual naming tasks, and one set of olfactory and visual matching tasks were administered to eight normal volunteers. In the olfactory naming task (ON), odors from familiar items, associated with some verbal label, were to be named. Hence, it required long-term olfactory memory retrieval for stimulus recognition. The olfactory matching task (OM) involved differentiating a recently encoded unfamiliar odor from a sequentially presented group of unfamiliar odors. This required short-term olfactory memory retrieval for stimulus differentiation. The simple olfactory and visual stimulation resulted in activation of the left orbitofrontal region, the right piriform cortex, and the bilateral occipital cortex. During olfactory naming, activation was detected in the left cuneus, the right anterior cingulate gyrus, the left insula, and the cerebellum bilaterally. It appears that the effort to identify the origin of an odor involved semantic analysis and some degree of mental imagery. During olfactory matching, activation was observed in the left cuneus and the cerebellum bilaterally. This identified the brain areas activated during differentiation of one unlabeled odor from the others. In cross-task analysis, the region found to be specific for olfactory naming was the left cuneus. Our results show definite recruitment of the visual cortex in ON and OM tasks, most likely related to imagery component of these tasks. The cerebellar role in cognitive tasks has been recognized, but this is the first PET study that suggests that the human cerebellum may have a role in cognitive olfactory processing as well.     

Puberty and olfactory preferences of males

The preferences of 302 males, aged 8–9, 14 and 16 years, for ten food related odors were determined using a hedonic scale. The aim was to establish if major changes in preferences occur during the period which encompasses puberty. Small but significant differences between the 8–9 year group and the others were obtained for meat and chicken odors; between the 14 year group and the others for peanut butter odor; and there was a change from dislike to like with age for coffee odor. The results were almost all confirmed in a retest. The general lack of differences between the responses of the three age groups indicated that no major change occurs in preferences for food odors between the ages of 8 and 16 years. Although these results indicate that olfactory preferences for food odors are not affected by puberty, they do not eliminate the possibility that preferences may be altered for odors related to sex and emotion [13].     

Dopamine D(2) receptor activation modulates perceived odor intensity

Dopaminergic modulation affects odor detection thresholds and olfactory discrimination capabilities in rats. The authors show that dopamine D(2) receptor modulation affects odor discrimination capabilities in a manner similar to the modulation of stimulus intensity. Performance in a simultaneous odor discrimination task was systematically altered by manipulations of both odorant concentration and D(2) receptor activation (agonist quinpirole, 0.025-0.5 mg/kg; antagonist spiperone, 0.5 mg/kg). Rats' discrimination performance systematically improved at higher odor concentrations. Blockade of D(2) receptors improved performance equivalent to increasing odor concentration by 2 log units, whereas activation of D(2) receptors reduced odor discrimination performance in a dose-dependent manner. Bulbar dopamine release may serve a gain control function in the olfactory system, optimizing its sensitivity to changes in the chemosensory environment.     

Olfactory-evoked regional cerebral blood flow in Alzheimer's disease

Olfaction is impaired in Alzheimer's disease (AD). It was hypothesized that AD would reduce olfactory-evoked perfusion in mesial temporal olfactory (piriform) cortex, where neuropathology begins. Seven AD patients and 8 elderly controls (ECs) underwent olfactory threshold and identification tests and olfactory stimulation during positron emission tomography. Odor identification was impaired in AD, but threshold was not. Olfactory stimulation in ECs activated right and left piriform areas and right anterior ventral temporal cortex. AD patients had less activation in right piriform and anterior ventral temporal cortex but not in the left piriform area. Although orbital cortex did not activate in ECs, there was a significant between-groups difference in this area. Right piriform activation correlated with odor identification. Impaired odor identification likely reflects sensory cortex dysfunction rather than cognitive impairment. Given olfactory bulb projections to the mesial temporal lobe, olfactory stimulation during functional imaging might detect early dysfunction in this region.     

Assessment of olfactory function and androstenone odor thresholds in humans with or without functional occlusion of the vomeronasal duct

To obtain information on the possible role of the vomeronasal duct (VND) in odor perception and human pheromone detection, the present study investigated different aspects of olfactory function, including thresholds for androstenone in adults with or without detectable VNDs. The study also examined correlations between detection thresholds of androstenone odor and general olfactory function. Subjects' olfaction was assessed with tests for odor identification, odor discrimination, and phenyl ethyl alcohol odor threshold. Measurements were performed on 1 side only, with and without covering the VND. Subjects with or without detectable VNDs did not differ in olfactory sensitivity or androstenone odor thresholds. A small but significant correlation was found between detection thresholds of androstenone and general olfactory function. Finally, covering of the VND did not affect olfactory function or androstenone sensitivity. Results suggest that the human VND does not play a major role in sensitivity toward odorants or the perception of androstenone.     

Olfaction in the psychosis prodrome: Electrophysiological and behavioral measures of odor detection

Smell identification deficits (SIDs) are relatively specific to schizophrenia and its negative symptoms, and may predict transition to psychosis in clinical high-risk (CHR) individuals. Moreover, event-related potentials (ERPs) to odors are reduced in schizophrenia. This study examined whether CHR patients show SIDs and abnormal olfactory N1 and P2 potentials. ERPs (49 channels) were recorded from 21 CHR and 20 healthy participants (13 males/group; ages 13–27 years) during an odor detection task using three concentrations of hydrogen sulfide (H₂S) or blank air presented unilaterally by a constant-flow olfactometer. Neuronal generator patterns underlying olfactory ERPs were identified and measured by principal components analysis (unrestricted Varimax) of reference-free current source densities (CSD). Replicating previous findings, CSD waveforms to H₂S stimuli were characterized by an early N1 sink (345 ms, lateral–temporal) and a late P2 source (600 ms, mid-frontocentroparietal). N1 and P2 varied monotonically with odor intensity (strong > medium > weak) and did not differ across groups. Patients and controls also showed comparable odor detection and had normal odor identification and thresholds (Sniffin' Sticks). However, olfactory ERPs strongly reflected differences in odor intensity and detection in controls, but these associations were substantially weaker in patients. Moreover, severity of negative symptoms in patients was associated with reduced olfactory ERPs and poorer odor detection, identification and thresholds. Three patients who developed psychosis had poorer odor detection and thresholds, and marked reductions of N1 and P2. Thus, despite the lack of overall group differences, olfactory measures may be of utility in predicting transition to psychosis among CHR patients.     

Enrichment to odors improves olfactory discrimination in adult rats

The authors tested how prior odor enrichment affects the spontaneous discrimination of both preexposed and novel odors. Experimental rats were exposed to single odors or to pairs of similar or dissimilar odors for 1-hr periods twice daily over 20 days. Spontaneous discriminations between pairs of similar odors were tested before and after the odor exposure period using an olfactory habituation task. The authors found that (a) experimental rats did not spontaneously discriminate similar odor pairs before the exposure period, whereas they spontaneously discriminated them after the enrichment period, and (b) the improvement of performance was not selective for the odors used during enrichment. These results show that odor experience changes perception in the manner predicted based on other groups' electrophysiological experiments.     

Differential effects of lesions of the vomeronasal and olfactory nerves on garter snake (Thamnophis sirtalis) responses to airborne chemical stimuli

The roles of the main (MOS) and accessory (AOS) olfactory systems of garter snakes in response to airborne chemicals were investigated. Preoperatively, all snakes responded to airborne odors with increased tongue-flick rate and duration. Postoperatively, sham-operated snakes responded to airborne odors with increased tongue-flick rates, but snakes with main olfactory nerve cuts failed to respond to the odors, and snakes with vomeronasal nerve cuts responded to nonprey odors only. Preoperatively, exposure to earthworm odor produced more frequent and shorter duration tongue-flicks during locomotion compared with exposure to water. Postoperatively, only sham-lesioned snakes exhibited differential responding to earthworm odors. This study demonstrates that the MOS is critical for the initiation of tongue-flick behavior in response to airborne odors and that discrimination of odors with biological significance requires a functional AOS.     

Distributed representation of social odors indicates parallel processing in the antennal lobe of ants

In colonies of eusocial Hymenoptera cooperation is organized through social odors, and particularly ants rely on a sophisticated odor communication system. Neuronal information about odors is represented in spatial activity patterns in the primary olfactory neuropile of the insect brain, the antennal lobe (AL), which is analog to the vertebrate olfactory bulb. The olfactory system is characterized by neuroanatomical compartmentalization, yet the functional significance of this organization is unclear. Using two-photon calcium imaging, we investigated the neuronal representation of multicomponent colony odors, which the ants assess to discriminate friends (nestmates) from foes (nonnestmates). In the carpenter ant Camponotus floridanus, colony odors elicited spatial activity patterns distributed across different AL compartments. Activity patterns in response to nestmate and nonnestmate colony odors were overlapping. This was expected since both consist of the same components at differing ratios. Colony odors change over time and the nervous system has to constantly adjust for this (template reformation). Measured activity patterns were variable, and variability was higher in response to repeated nestmate than to repeated nonnestmate colony odor stimulation. Variable activity patterns may indicate neuronal plasticity within the olfactory system, which is necessary for template reformation. Our results indicate that information about colony odors is processed in parallel in different neuroanatomical compartments, using the computational power of the whole AL network. Parallel processing might be advantageous, allowing reliable discrimination of highly complex social odors.