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Comparison of models of odor interaction.

Subjects rated the overall perceived intensity of concentrations of the odorants cineole, geraniol, hexyl salicylate, and linalyl acetate smelled alone and in binary mixtures. The subjects also rated intensity of specified constituents (e.g. amount of cineole in cineole, and in mixtures of cineole and linalyl acetate). The intensity of the stronger component alone offered a close description of perceived intensity. In addition to the Stronger Component model, two other psychological models (Vector and U model) and two psychophysical models (UPL2 and Equiratio Mixture model) offered descriptions ranging from fair to very good. Psychological models gave better fits, but lack explanatory power. Some results indicated that weaker odors add more potently than stronger odors, an outcome incompatible with these models. The psychophysical models, based on the additivity of single components, generally overestimated perceived intensity. Judgments of individual qualities gave only slight encouragement to any expectation of differences in masking or maskability among odorants. The results highlight the need to test particular critical hypotheses regarding how people perceive mixtures.

Acyclic Monoterpenes↗

Cognitive processes in odorant mixture assessment.

'Comparison of models of odor interaction' by Cain and his colleagues (1995) is a turning point in assessment of the strengths of smell of mixtures of odorants. Their paper provides the best empirical comparison to date of models of (olfactory) binary mixture intensities and combines this analysis with a cogent critique of the whole tradition of mixture psychophysics. This commentary on their paper argues that their critique is not radical enough.

Cognition↗

Is short-term odour recognition predictable from odour profile?

Smelling an odour induces a pattern of sensations, images and memories which participate in identification. It was proposed that perceptual memory performances for odours could be inferred from the description of these olfactory representations. The subject was asked to elaborate an odour descriptor profile, and a short-term odour recognition memory task was chosen to test the individual perceptual memory performance. Two pattern-recognition methods based on artificial neural networks and discriminant analysis were carried out and allowed odour profile and perceptual memory performance to be related. Insofar as the subjects gave dichotomic responses in the recognition memory task, each response could be evaluated in terms of correct or incorrect responses. Simulations indicated that the olfactory recognition memory performance can be predicted in man from odour-elicited semantic profiles by using artificial neural networks. It was also shown that all semantic descriptors do not participate in olfactory recognition to the same degree. Low-level information, such as intensity, familiarity and hedonic judgements, did not allow the artificial neural network to predict the olfactory performance. By contrast, high-level information, such as gustatory, olfactory and visual evocations, allowed artificial networks to make such predictions.

Adult↗

Olfactory evoked potential produced by electrical stimulation of the human olfactory mucosa.

Most physiological studies of the human olfactory system have concentrated on the cortical level; the olfactory bulbar level has been studied rarely. We attempted to stimulate the human olfactory mucosa by electrical pulse to detect the bulbar potentials. Electrical stimulation (2 mA, 0.5 ms) of the human olfactory mucosa evoked a change in potential recorded from the frontal sector of the head. A negative peak of the evoked potential that occurred at 19.4 ms (grand means, n = 5) after stimulation was the clearest. The highest amplitude of the potential was recorded from the frontal sector of the head on the stimulated side. Our findings were similar to the experimental results obtained from the olfactory bulbs of animals. This evoked potential was considered to be the human olfactory bulbar potential. When the subjects were stimulated by applying electricity to the olfactory mucosa, no sensation of smell occurred even though evoked potentials were recorded. Evoked potentials were recorded only when the stimulating electrode was located in the olfactory cleft. When the stimulating electrode was outside the olfactory cleft, the stimulation caused pain. The trigeminal nerve seemed to be stimulated by electricity. Olfactory evoked potentials produced by the electrical stimulation of the human olfactory mucosa should aid the research on human olfactory physiology, and may be applicable to clinical tests of olfactory dysfunction.

Electric Stimulation↗

Odor hedonics: connection with emotional response estimated by autonomic parameters.

The aim of this paper is to analyse the relationship between self-report hedonic evaluations and the physiological expression of emotion in response to odorants. We try to solve the following questions: (1) Is it possible to find any experimental evidence that the sense of smell is linked with emotion? (2) What kind of odorants can be distinguished by autonomic analysis? (3) Is there a link between hedonics and autonomic information? The effects of odorants on the emotional process were estimated, in terms of autonomic nervous system (ANS) activity. Fifteen subjects inhaled five odorants as olfactory stimuli: lavender (LAV), ethyl acetoacetate (EAA), camphor (CAM), acetic acid (AA) and butyric acid (BA). After inhaling the odorant, subjects were requested to fill out an 11-point hedonic scale to rate its pleasantness versus unpleasantness. ANS parameters were as follows: two electrodermal responses, skin potential (SP) and resistance (SR); two thermovascular parameters, skin blood flow (SBF) and skin temperature (ST); and two cardiorespiratory parameters; instantaneous respiratory frequency (IRF) and instantaneous heart rate (IHR). Simultaneous recording of six parameters showed that specific autonomic patterns were associated with each odorant. An analysis of variance made it possible to differentiate among the five odorants. Two-by-two odorant comparisons for autonomic responses using Tukey's HSD multiple comparison test only permitted differentiation between pleasant odorants (LAV and EAA) and unpleasant (AA and BA) ones, but camphor was differentiated from both pleasant and unpleasant odorants. Each odorant elicited responses in the different parameters, yet subjects responded through their preferential channels; an average of two channels was used by each subject. These results when compared with those obtained with other senses (visual and auditory), did not evidence the postulated preferential link between olfaction and emotion. A strong link between hedonics and ANS response could be demonstrated when considering each subject and mainly through his/her preferential channel(s); conversely a weak correlation (SR duration excepted) was obtained between inter-subjects' hedonic evaluation. It seems that for a given population the autonomic response reflect the odor valence only through some parameters related to the main preferential channel(s) and thus the global autonomic pattern has to be considered.

Acetic Acid↗

Decline in taste and odor discrimination abilities with age, and relationship between gustation and olfaction.

It is important to learn about changes in both taste and odor perceptions with increasing age, because the taste of foods we encounter in our daily life is strongly affected by their smell. This study discusses the difference in qualitative taste and odor discrimination between the elderly and the young. Tastants and odorants used in this study were presented not as single stimuli but as a taste mixture (sucrose and tartaric acid) and an odor mixture (beta-phenylethyl alcohol and gamma-undecalactone). The results showed that quality discrimination abilities of the elderly subjects for both taste and odor were significantly lower than those of the young subjects, indicating a decline in quality discrimination abilities related to age. Also, a moderate but significant correlation was observed between the taste discrimination ability and the odor discrimination ability. We measured thresholds for single-taste and odor components in mixtures and compared them between the elderly and the young to investigate the cause for these findings.

Adult↗

Quantification of odor quality.

The relationship between odor quality and molecular properties is arguably the most important issue in olfaction. Despite sophistication in the chemical characterization of molecules, accompanying perceptual characterization has had little quantitative usefulness, relying mostly on enumerative description. As a result of weak interest in the topic outside industry and little agreement regarding how to measure quality, the field of olfactory psychophysics has failed to develop a substantial database for odor quality and has offered little help to other researchers, e.g. neurobiologists, in choice of stimuli, interpretation of outcome or testable hypotheses. This review scrutinizes how psychophysicists and others have measured quality and offers criteria for useful techniques. Most measures have had a subjective component that makes them anachronistic with modern methodology in experimental behavioral science, indeterminate regarding the extent of individual differences, unusable with infrahumans and of unproved ability to discern small differences. Techniques based upon performance, rather than on the more common reporting of mental content, offer firmer possibilities for growth. These techniques inevitably tap the discriminative basis of perception. The nonsubjective techniques have high sensitivity, can have counterparts in infrahuman research, are suitable to examine individual differences and yield non-negotiable answers with potential archival value. Discriminative techniques have their limitations, too-principally excess sensitivity that abridges their use to comparisons between similar-smelling stimuli. Research has begun to extend that range and may overcome the limitation. Application of discriminative methods may have the side-effect of shifting focus in structure-activity research from searches for molecular least common denominators that underlie often vague similarity to the search for molecular properties of importance in discrimination of small differences.

Humans↗

Functional genomics of odor-guided behavior in Drosophila melanogaster.

The avoidance response to repellent odorants in Drosophila melanogaster, a response essential for survival, provides an advantageous model for studies on the genetic architecture of olfactory behavior. Transposon tagging in a highly inbred strain of flies in combination with a rapid and simple statistical behavioral assay enables the identification of not only large phenotypic effects, but also small aberrations from wild-type avoidance behavior. The recent completion of the sequence of the Drosophila genome facilitates the molecular characterization of transposon-tagged genes and correlation between gene expression and behavior in smell-impaired (smi) mutant lines. Quantitative genetic analyses of a collection of smi lines in a co-isogenic background revealed an extensive network of epistatic interactions among genes that shape the olfactory avoidance response. Candidate genes for several of these transposon-tagged smi loci implicate genes that mediate odorant recognition, including a novel odorant binding protein; signal propagation, including a voltage-gated sodium channel; and a protein containing multiple leucine rich repeats and PDZ domains likely to be involved in postsynaptic organization in the olfactory pathway. Several novel genes of unknown function have also been implicated, including a novel tyrosine-regulated protein kinase. The discovery and characterization of novel gene products that have major, hitherto unappreciated effects on olfactory behavior will provide new insights in the generation and regulation of odor-guided behavior. The identification and functional characterization of proteins encoded by smi genes that form part of the olfactory subgenome and correlation of polymorphisms in these genes with variation in odor-guided behavior in natural populations will advance our understanding of the genetic architecture of chemosensory behavior.

Animals↗

The influence of olfactory concept on the probability of detecting sub- and peri-threshold components in a mixture of odorants.

The headspace of apple juice was analysed to obtain an ecologically relevant stimulus model mixture of apple volatiles. Two sets of volatiles were made up: a set of eight supra-threshold volatiles (MIX) and a set of three sub-threshold volatiles. These sets were used to test the hypothesis that sub-threshold components can change the quality of a familiar smelling mixture of odorants when added to this mixture. In order to test this hypothesis, three successive dilutions of the sub-threshold volatiles were prepared in such a way that the strongest was at the threshold concentration and the two lower concentrations were below the threshold. The detection probabilities of the sub-threshold components in a blank stimulus were compared with the detectabilities in MIX. The sub- and peri-threshold volatiles were detected no better in MIX than in a blank. On the contrary, sub- and peri-threshold volatiles were better detected alone than when added to MIX. However, when the group of subjects was split into two sub-groups, employing either a rough or a detailed concept definition of the target stimulus, respectively, the subjects with highly refined concepts were better able to detect the presence of sub-threshold volatiles in MIX than those with poorly refined stimulus concepts. The effect of stimulus concept definition occurred independently of the proportions of correct detections of sub-threshold volatiles in a blank.

Adolescent↗

Distortion of olfactory perception: diagnosis and treatment.

Clinically, olfaction can fail in any of three ways: (i) decreased sensitivity (hyposmia, anosmia) and two types of distortion (dysosmia); (ii) distorted quality of an odorant stimulation (troposmia); (iii) perceived odor when no odorant is present (phantosmia, hallucination). The distortions are usually much more upsetting to a person's quality of life than a simple loss. An ipsilatersal loss of olfactory sensitivity is often identified in the nostril with any type of olfactory distortion. The pathophysiology of a stimulated distortion (troposmia) is likely a decreased number of functioning olfactory primary neurons so that an incomplete characterization of the odorant is made. In phantosmia, two possible causations include an abnormal signal or inhibition from the primary olfactory neurons or peripheral olfactory or trigeminal signals that "trigger" a central process. The clinician's goal is to carefully define the problem (e.g. taste versus smell, real versus perceived, one versus two nostrils), to perform the appropriate examination and testing and to provide therapy if possible. Treatment includes assurance with no active therapy (because many of these will naturally resolve), topical medications, systemic medications, anesthesia to parts of the nose and, rarely, referral for surgical excision of olfactory neurons. Endoscopic transnasal operations have the advantage of treating phantosmia and sometimes allowing a return of olfactory ability after the operation.

Humans↗

Autonomic nervous system responses to odours: the role of pleasantness and arousal.

Perception of odours can provoke explicit reactions such as judgements of intensity or pleasantness, and implicit output such as skin conductance or heart rate variations. The main purpose of the present experiment was to ascertain: (i) the correlation between odour ratings (intensity, arousal, pleasantness and familiarity) and activation of the autonomic nervous system, and (ii) the inter-correlation between self-report ratings on intensity, arousal, pleasantness and familiarity dimensions in odour perception. Twelve healthy volunteers were tested in two separate sessions. Firstly, subjects were instructed to smell six odorants (isovaleric acid, thiophenol, pyridine, L-menthol, isoamyl acetate, and 1-8 cineole), while skin conductance and heart rate variations were being measured. During this phase, participants were not asked to give any judgement about the odorants. Secondly, subjects were instructed to rate the odorants on dimensions of intensity, pleasantness, arousal and familiarity (self-report ratings), by giving a mark between 1 (not at all intense, arousing, pleasant or familiar) and 9 (extremely intense, arousing, pleasant or familiar). Results indicated: (i) a pleasantness factor correlated with heart rate variations, (ii) an arousal factor correlated with skin conductance variations, and (iii) a strong correlation between the arousal and intensity dimensions. In conclusion, given that these correlations are also found in other studies using visual and auditory stimuli, these findings provide preliminary information suggesting that autonomic variations in response to olfactory stimuli are probably not modality specific, and may be organized along two main dimensions of pleasantness and arousal, at least for the parameters considered (i.e. heart rate and skin conductance).

Adult↗

Frontalin: a chemical message of musth in Asian elephants (Elephas maximus).

Musth is an important male phenomenon affecting many aspects of elephant society including reproduction. During musth, the temporal gland secretions (as well as the urine and breath) of adult male Asian elephants (Elephas maximus) discharge a variety of malodorous compounds together with the bicyclic ketal, frontalin. In contrast, teenage male elephants in musth release a sweet-smelling exudate from their facial temporal gland. We recently demonstrated that the concentration of frontalin becomes increasingly evident as male elephants mature. In the present study, we demonstrate that behaviors exhibited towards frontalin are consistent and dependent on the sex, developmental stage and physiological status of the responding conspecific individual. To examine whether frontalin functions as a chemical signal, perhaps even a pheromone, we bioassayed older and younger adult males, and luteal- and follicular-phase and pregnant females for their chemosensory and behavioral responses to frontalin. Adult males were mostly indifferent to frontalin, whereas subadult males were highly reactive, often exhibiting repulsion or avoidance. Female chemosensory responses to frontalin varied with hormonal state. Females in the luteal phase demonstrated low frequencies of responses, whereas pregnant females responded significantly more frequently, with varied types of responses including those to the palatal pits. Females in the follicular phase were the most responsive and often demonstrated mating-related behaviors subsequent to high chemosensory responses to frontalin. Our evidence strongly suggests that frontalin, a well-studied pheromone in insects, also functions as a pheromone in the Asian elephant: it exhibits all of the determinants that define a pheromone and evidently conveys some of the messages underlying the phenomenon of musth.

Aging↗

Ratings of overall olfactory function.

The aim of this study was to investigate the accuracy of self-reported ratings of olfactory function in 83 healthy subjects. Such ratings were compared with quantitative measures of olfactory function, as well as with ratings of nasal patency. In experiment 1 subjects rated olfactory function and nasal patency before olfactory testing, whereas in experiment 2 the reverse was the case. No feedback regarding test results were provided until after completion of the testing. The principal findings were: (i) when ratings preceded measurements of olfactory function, there was no significant correlation between the two parameters. However, ratings of olfactory function correlated significantly with ratings of nasal airway patency. (ii) In contrast, when measurements of olfactory function preceded the ratings, this constellation switched. Now ratings of olfactory function correlated significantly with measured olfactory function, whereas there was no significant correlation between ratings of nasal airway patency and ratings of olfactory function. In conclusion, these data suggest that ratings of olfactory function are unreliable in healthy, untrained subjects. The ratings seem to reflect changes of nasal airway patency to a larger degree than measurable olfactory function. The results further indicate that this is mainly due to the limited attention the sense of smell receives in daily life.

Adult↗

Human odor detectability: new methodology used to determine threshold and variation.

Current ambiguity concerning the related issues of optimal means for measurement of odor sensitivity and the functional properties of the olfactory system hinders progress in basic and applied research on the human sense of smell. To address these needs, we selected n-amyl acetate (nAA) as a test odorant and developed a methodology in which participants (Ps) receive multiple presentations each session of several concentrations. Yes-no responses as to whether odor was detected are analyzed using binomial statistics, with the probability that a given proportion of yes responses (or greater) would occur by chance alone being treated as the inverse of detectability. Over the course of multiple sessions, this information is also used to maximize the collection of data in the peri-threshold region. Surprisingly, data collected over as many as 14 sessions were fit well by a single logistic regression model relating probability and concentration. Threshold concentrations, defined as those corresponding to a probability of 0.05, varied from 7.11 to 167.53 p.p.b. (v/v) for 11 Ps. Our approach and findings, if shown to be representative of other combinations of Ps and odorants, could accelerate the pace of research in human olfaction by providing a comprehensive operational definition of the limit of the olfactory system to detect odorant molecules.

Adult↗

Hemodynamic response of the frontal cortex elicited by intravenous thiamine propyldisulphide administration.

The intravenous olfaction (IVO) test is a unique type of clinical olfactometry and is widely used in Japan. However, it is difficult to distinguish actual olfactory disturbance from feigned disturbance because the IVO test is a psychophysical test. To resolve this problem, we investigated the possibility of an objective IVO test assisted with near infrared spectroscopy (NIRS). IVO testing was performed according to the usual protocol with thiamine propyldisulphide (alinamin) administration. The relative oxy- and deoxyhemoglobin levels of the orbitofrontal area during olfactory stimulation by IVO test were measured by NIRS. Pairs of NIRS emitters and detectors were positioned on the bilateral frontal scalp. After administration of alinamin, oxyhemoglobin levels increased, though deoxyhemoglobin levels did not change. An increase in oxyhemoglobin levels was observed bilaterally. Administration of saline did not elicit any change in the oxy- or deoxyhemoglobin levels and concentration of the administered alinamin related increasing of the oxyhemoglobin level was observed. Oxyhemoglobin remained unchanged in anosmic subjects despite administration of alinamin. The latency of oxyhemoglobin increase on each side and smelling latency showed significant correlation. Latencies of oxyhemoglobin increases between the right and left sides also showed significant correlation. Oxyhemoglobin response appears to be linked to olfactory related response. NIRS is a useful technique for the development of an objective form of IVO testing.

Adult↗

The effect of emotion and personality on olfactory perception.

It is well established that both the emotional tone of sensory stimuli and the personality characteristics of an individual can bias sensory perception. What has largely been unexplored is whether the current emotional state of an individual has a similar effect, and how it works together with other factors. Here we carry out a comprehensive study to examine how olfactory perception is affected by the emotional tone of the stimuli, and the personality and current emotional state of the individual. Subjects reported experiencing happiness, sadness, negativity/hostility and neutrality when exposed to corresponding emotionally themed video clips, and in each case, smelled a suprathreshold pleasant, an unpleasant and a neutral odorant. The time taken for the subject to detect each odorant and the olfactory intensity were recorded. We found that women detected the pleasant odorant faster than the neutral one. In addition, personality modulated reaction time and olfactory intensity, such that neurotic and anxious individuals were selectively biased toward affective rather than neutral odorants. Finally, current emotional state augmented intensity in men but not in women, and differentially influenced the response time. These findings provided new insights into the effects of emotion and personality on olfactory perception.

Adult↗

A metamemory perspective on odor naming and identification.

A metacognitive perspective is utilized to elucidate why it is so difficult to name common odors and what characterizes the subjective knowledge people have about their actual odor knowledge. Odor-naming failures are often accompanied by strong feelings of knowing (FOK) or feelings of imminent retrieval of what it is that smells. The paper's two experiments investigate FOK judgements and tip of the tongue (TOT) experiences for odor and person names. The data indicate that our inability to correctly name odors are typically not due to the often proposed uniquely poor association between odors and their proper names, but rather due to failures to identify the odors, that is, failures to know 'what it is'. It was also found that (i) TOT experiences are very unusual for odor names and more so than for person names; (ii) FOK judgements about odor names are significantly less predictive of later retrieval than equivalent judgements about names of persons; (iii) FOK judgements were highly correlated with the familiarity of the cue (odor or picture of famous person), rendering some support for the idea that FOK judgements are based on the perceived familiarity of the cue triggering the FOK; and (iv) the idea that FOK judgements are based on the amount of available information about the sought-for memory (accessibility theory) was also supported.

Adult↗

Cross-modal interactions between olfaction and vision when grasping.

This study used kinematics to investigate the integration between vision and olfaction during grasping movements. Participants were requested to smell an odorant and then grasp an object presented in central vision. The results indicate that if the target was small (e.g., a strawberry), the time and amplitude of maximum hand aperture were later and greater, respectively, when the odor evoked a larger object (e.g., an orange) than when the odor evoked an object of a similar size as the target or no odor was presented. Conversely, the time and amplitude of maximum hand aperture were earlier and reduced, respectively, when the target was large (e.g., a peach) and the odor evoked a smaller sized object (e.g., an almond) than when the odor evoked an object of a similar size as the target or no odor was presented. Taken together, these results support the evidence of cross-modal links between olfaction and vision and extend this notion to goal-directed actions.

Adult↗