Perception of pungent odorants alone and in binary mixtures

The present investigation explores the stimulus–response(psychophysical) functions for total nasal perceived intensityfor two pungent odorants, formaldehyde and ammonia, presentedeither alone or with different backgrounds of the other irritant.Stimuli were comprised of four formaldehyde concentrations (1.0,3.5, 6.9 and 16.7 p.p.m.); four ammonia concentrations (210,776, 1172 and 1716 p.p.m.); and their 16 binary mixtures. Ammoniafunctions showed a consistent upward concavity. At low, mediumand high concentrations, the total perceived intensity of themixtures showed hypoadditivity, simple additivity and hyperadditivity,respectively. That is, the intensity of the corresponding mixtureswas significantly lower than, equal to, or greater than thesum of its components. The progressive involvement of pungency,aroused by common chemical sense stimulation, may be responsiblefor the increasing additivity observed. Simple additivity, andeven hyperadditivity, may characterize mixtures involving commonchemical stimulation.     

Enhancement of retronasal odors by taste

Psychophysical studies of interactions between retronasal olfaction and taste have focused most often on the enhancement of tastes by odors, which has been attributed primarily to a response bias (i.e., halo dumping). Based upon preliminary evidence that retronasal odors could also be enhanced by taste, the present study measured both forms of enhancement using appropriate response categories. In the first experiment, subjects rated taste ("sweet," "sour," "salty," and "bitter") and odor ("other") intensity for aqueous samples of 3 tastants (sucrose, NaCl, and citric acid) and 3 odorants (vanillin, citral, and furaneol), both alone and in taste-odor mixtures. The results showed that sucrose, but not the other taste stimuli, significantly increased the perceived intensity of all 3 odors. Enhancement of tastes by odors was inconsistent and generally weaker than enhancement of odors by sucrose. A second experiment used a flavored beverage and a custard dessert to test whether the findings from the first experiment would hold for the perception of actual foods. Adding sucrose significantly enhanced the intensity of "cherry" and "vanilla" flavors, whereas adding vanillin did not significantly enhance the intensity of sweetness. It is proposed that enhancement of retronasal odors by a sweet stimulus results from an adaptive sensory mechanism that serves to increase the salience of the flavor of nutritive foods.     

Perceptual interactions in odour mixtures: odour quality in binary mixtures of woody and fruity wine odorants

The qualitative perceptual interactions in three binary mixtures of wine odorants were studied: isoamyl acetate (fruity note)/whisky lactone (woody note), ethyl butyrate (fruity note)/whisky lactone (woody note) and ethyl butyrate (fruity note)/guaiacol (woody note). For each binary mixture, the perceived quality and intensity of 24 stimuli (four supra-threshold concentration levels of each of the two compounds and their 16 possible combinations) were evaluated in five replications by a trained panel of 13 subjects. The application of the Olsson predictive model for odour intensity and quality perception gave quite a good estimation of the evolution of single component identification in the mixture when the intensity proportion of unmixed components varied. However, this model was unable to account for the odour quality dominance in mixtures of iso-intense components. An alternative linear logistic model was proposed to study the qualitative dominance of the woody note in the three mixtures when the perceived intensities of each unmixed compound were equal.     

Age-related deficiency in the perceived strength of six odorants

A group of 20 elderly persons (70–89 yr) and a controlgroup of 20 young persons (18–25 yr) made magnitude estimationsof five concentration levels of six odorants and of five concentrationlevels of a tastant, NaCl. Relative to the estimations of thesalt solutions, the elderly's estimations of all six odorantswere lower than those of the young. This outcome substantiatesan earlier finding that, at least for one odorant (iso-amylbutyrate), old age blunts perceived odor strength more frequentlyand seriously than gustatory strength. The present experimentbroadens the picture and leads to the conclusion that age-relatedhyposmia is likely to affect the perception of many, if notall, odors. The six odorants were selected on the basis of structuraldiversity, hedonic tone, and earlier psychophysical study ofthem. They include three pleasant odors (iso-amyl butyrate,benzaldehyde, and d-limonene), one foul smelling (pyridine),and two relatively neutral ones (ethyl and iso-amyl alcohol).To a first approximation age-related weaknesses to these compoundscan be characterized as a constant per cent reduction of olfactorystrength across concentration level. tion level.     

Computer modeling of information processing mechanisms in the olfactory system. III. Reproduction of psychophysical phenomena by the olfactory bulb model

Psychophysical phenomena typical of olfaction were reproduced using a computer model of olfactory bulb. The procedure of numerical experiments is described. The model reproduces the following phenomena: fusion of odors, strong and weak odors, suppression of weak odors by strong odors, indemnity of odors, changes in odor with time, consecutive olfactory images, sensibilization, consecutive olfactory constrast, and synergims. It was concluded that computer-assisted experimentation in combination with neurophysiological and psychophysical experiments can considerably increase the efficiency of research of odorants and the olfaction process.     

Just noticeable differences in component concentrations modify the odor quality of a blending mixture

The odors we perceive are mainly the result of mixtures of odorants that, however, are commonly perceived as single undivided entities; nevertheless, the processes involved remain poorly explored. It has been recently reported that perceptual blending based on configural olfactory processing can cause odorant mixtures to give rise to an emergent odor not present in the components. The present study examined whether specific component proportions are required to elicit an emergent odor. Starting from the composition of a ternary target mixture in which an emergent pineapple odor was perceived, 4 concentration levels of each component were chosen to elicit just noticeable differences (JNDs). Each combination of levels was used to design sample mixtures. Fifteen subjects evaluated the intensity, typicality, and pleasantness of each sample mixture against the target mixture in a paired-comparison protocol. Statistical modeling showed that a variation of less than 1 JND in one of the components was sufficient to induce a significant decrease in pineapple odor typicality in the ternary mixture. This finding confirms previous findings on perceptual blending in simple odorant mixtures and underscores the human ability to discriminate between odor percepts induced by mixtures including very similar odorant proportions.     

Newborn Rabbit Perception of 6-Odorant Mixtures Depends on Configural Processing and Number of Familiar Elements

Perception of odors, i.e. usually of mixtures of odorants, is elemental (the odorants'' odor qualities are perceived in the mixture) or configural (the odor quality of the mixture differs from the one of each odorant). In human adults, the Red Cordial (RC) mixture is a configurally-processed, 6-odorant mixture. It evokes a red cordial odor quality while none of the elements carries that odor. Interestingly, in newborn rabbits, the same RC mixture is weak configurally perceived: the newborns behaviorally respond to all the elements after conditioning to the whole mixture, but not to the mixture after conditioning to a single element. Thus, they perceive in the RC mixture both the odor quality of the RC configuration and the quality of each element. Here, we aimed to determine whether this perception is modulated by quantitative (number of elements) and/or qualitative bits of information (nature of elements) previously learned by the animals. Newborns were conditioned to RC sub-mixtures of different complexity and composition before behavioral testing to RC. Pups generalized their sucking-related response to RC after learning at least 4 odorants. In contrast, after conditioning to sub-mixtures of another 6-odorant mixture, the elementally perceived M^V mixture, pups responded to M^V after learning one or two odorants. The different generalization to RC and M^V mixtures after learning some of their elements is discussed according to three hypotheses: i) the configural perception of RC sub-mixtures, ii) the ratio of familiar/unfamiliar individual information elementally and configurally perceived, iii) the perception of RC becoming purely elemental. The results allow the first hypothesis to be dismissed, while further experiments are required to distinguish between the remaining two.     

Color enhances orthonasal olfactory intensity and reduces retronasal olfactory intensity

The effect of color on orthonasal and retronasal odor intensities was investigated. When odorants were smelled orthonasally (i.e., through the nostrils), color enhanced odor intensity ratings, consistent with previous reports. However, when odorants were smelled retronasally (i.e., the odorous solution was put in the mouth), color reduced odor intensity ratings. These different effects of color on odor intensity (i.e., enhancement orthonasally and suppression retronasally) appear to be the result of route of olfactory stimulation rather than of any procedural artifact. This supports previous reports that retronasal and orthonasal odors are perceived differently.     

Effects of perceived and imagined odors on taste detection

We assessed the influence of different odors on detection of a sweet tastant, and the ability of imagined odors to elicit the same effects as perceived odors on taste perception. The tastant used was sucrose, and the two odorants were strawberry and ham. In the first experiment, participants either smelled or imagined one of two odors during taste detection tasks (between-subject design), whereas in the second one, subjects completed both the odor imagery and perception conditions with taste detection tasks (within-subject design). The effect was odorant-specific: detection of sucrose was significantly better when subjects smelled strawberry than when they smelled ham. Furthermore, imagined odors influenced taste perception in the same way as did perceived odors. We concluded that the odor-specific effect on taste perception is an authentic perceptual phenomenon. Our results also support the notion that odor-induced changes in taste perception are mediated centrally. Finally, our findings are in agreement with reports supporting the existence of odor imagery.     

Response of olfactory receptor neurons in honeybees to odorants and their binary mixtures

Recordings were made from single sensilla placodea of the worker honeybee (Apis mellifera). The sensilla were stimulated with one of two sets of four compounds and their binary mixtures, at two dosage levels. Aromatic compounds comprised one set, and saturated n-octane derivatives comprised the other set. Correlation, principal component, and cluster analyses indicate that responses to binary mixtures are not linear combinations of responses to the component compounds. The first principal component indicated that neuronal units had either more excitatory or more inhibitory responses to all odorants than would be expected from a model where inhibitory and excitatory responses are randomly distributed among the neuronal units. When compared to the responses to the component odorants, synergistic responses to binary odors occurred more often than would be expected by chance. Clear inhibitory responses to binary odors were less prevalent. This study agrees with an earlier study employing aromatic odorants in that most of the aromatic odorants each had groups of receptor neurons that were relatively selective for it, and each odorant had a distinctly different number of receptor neurons selective for it. Among the octane derivatives, receptor neurons were selective for the level of oxidation of the functional group or its site of attachment, rather than specific compounds.     

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     

Odor representations in the rat olfactory bulb change smoothly with morphing stimuli

Many species of mammals are very good at categorizing odors. One model for how this is achieved involves the formation of "attractor" states in the olfactory processing pathway, which converge to stable representations for the odor. We analyzed the responses of rat olfactory bulb mitral/tufted (M/T) cells using stimuli "morphing" from one odor to another through intermediate mixtures. We then developed a phenomenological model for the representation of odors and mixtures by M/T cells and show that >80% of odorant responses to different concentrations and mixtures can be expressed in terms of smoothly summing responses to air and the two pure odorants. Furthermore, the model successfully predicts M/T cell responses to odor mixtures when respiration dependence is eliminated. Thus, odor mixtures are represented in the bulb through summation of components, rather than distinct attractor states. We suggest that our olfactory coding model captures many aspects of single and mixed odor representation in M/T cells.     

From molecule to mind: the role of experience in shaping olfactory function

The olfactory system is faced with a particular problem – the high dimensionality and inherent unpredictability of the chemical world. Most natural odorants encountered in everyday life are complex mixtures of many different volatiles. This means that from the outset the olfactory system has to contend with a great and often unpredictable diversity of molecules, making it difficult for stable primary features of the chemical world to be mapped onto the sensory surface. One solution to such unpredictability is provided by learning. Learning confers flexibility, enabling individuals of a given species to acquire and make use of the most appropriate information in a particular environment. Two examples of this are presented: learning of maternal odors in neonatal rabbits, including evidence that the sensory surface itself may be influenced by environmental conditions so as to enhance sensitivity to molecules of particular ecological relevance, and cross-cultural human studies suggesting that experience with everyday odors influences not only the way these are evaluated, but also their perceived intensity. It is concluded that an adequate understanding of odor coding and olfactory function will not be possible without taking such experience-dependent factors into account. Accepted: 11 May 1999     

Emotional processing of odors: evidence for a nonlinear relation between pleasantness and familiarity evaluations

Pleasantness, familiarity, and intensity are 3 interdependent dimensions commonly used to describe the perceived qualities of an odor. In particular, many empirical studies have demonstrated a positive correlation between familiarity and pleasantness. However, on the basis of both theoretical and methodological perspectives, we questioned the validity of such a relation for malodors. We report 2 studies based on subjective judgments of a large sample of odorants (Experiment 1) associated with autonomic recordings (Experiment 2). Multivariate exploratory analysis performed on the data splits the whole odorant set into 2 subsets composed, respectively, of unpleasant and pleasant odorants. Subsequent correlation analyses have shown that the relation between pleasantness and familiarity is specific for the pleasant odors in the 2 experiments. Moreover, autonomic activity was more important in response to malodors than to pleasant odors and was significantly correlated with unpleasantness ratings in the subset of unpleasant odors. These 2 studies argue in favor of a functional dissociation in the relations between both subjective and autonomic responses to odors as a function of pleasantness and indicate that researchers in the olfactory domain should consider the relations between pleasantness and familiarity as more complex than linear.     

Combinatorial receptor codes for odors

The discriminatory capacity of the mammalian olfactory system is such that thousands of volatile chemicals are perceived as having distinct odors. Here we used a combination of calcium imaging and single-cell RT-PCR to identify odorant receptors (ORs) for odorants with related structures but varied odors. We found that one OR recognizes multiple odorants and that one odorant is recognized by multiple ORs, but that different odorants are recognized by different combinations of ORs. Thus, the olfactory system uses a combinatorial receptor coding scheme to encode odor identities. Our studies also indicate that slight alterations in an odorant, or a change in its concentration, can change its "code," potentially explaining how such changes can alter perceived odor quality.     

Peripheral and central olfactory tuning in a moth

Animals can be innately attracted to certain odorants. Because these attractants are particularly salient, they might be expected to induce relatively strong responses throughout the olfactory pathway, helping animals detect the most relevant odors but limiting flexibility to respond to other odors. Alternatively, specific neural wiring might link innately preferred odors to appropriate behaviors without a need for intensity biases. How nonpheromonal attractants are processed by the general olfactory system remains largely unknown. In the moth Manduca sexta, we studied this with a set of innately preferred host plant odors and other, neutral odors. Electroantennogram recordings showed that, as a population, olfactory receptor neurons (ORNs) did not respond with greater intensity to host plant odors, and further local field potential recordings showed that no specific amplification of signals induced by host plant odors occurred between the first olfactory center and the second. Moreover, when odorants were mutually diluted to elicit equally intense output from the ORNs, moths were able to learn to associate all tested odorants equally well with food reward. Together, these results suggest that, although nonpheromonal host plant odors activate broadly distributed responses, they may be linked to attractive behaviors mainly through specific wiring in the brain.     

Potential mechanisms of retronasal odor referral to the mouth

The current study took a first step toward elucidating the sensory input that drives retronasal odor referral to the mouth. In 2 experiments, subjects performed odor localization tasks under various oral-nasal stimulation conditions that allowed us to assess the effects of direction of airflow, taste, and tactile stimulation on retronasal odor referral. Subjects reported the locations of perceived odors when food odorants were inhaled through the mouth alone or in the presence of water or various tastants in the mouth. The results indicated that when perceived alone, vanilla and soy sauce odor were localized 54.7%: 26.4%: 18.9% and 60.0%: 21.7%: 18.3% in the nose, oral cavity, and on the tongue, respectively. The localization of odors alone was not significantly different from when water was presented simultaneously in the mouth, indicating that tactile stimulation itself is not sufficient to enhance odor referral. However, the presence of sucrose, but not other tastes, significantly increased localization of vanilla to the tongue. Likewise, only NaCl significantly augmented referral of soy sauce odor to the tongue. These data indicate that referral of retronasal odors to the mouth can occur in the absence of a either taste or touch but that referral to the tongue depends strongly on the presence of a congruent taste.     

Sensitivity to odours in the embryo of the domestic fowl

Sensitivity to odours in the embryo of the domestic fowl was investigated on the day before hatching. Embryos were tested with four odorants: dichloroethane, cineole, amyl acetate and formic acid. Three odorants (dichloroethane, formic acid and cineole) produced an increase in the heart rate and a rise in the rates of beak-clapping and the first two increased the amount of head-shaking. Odorants had little effect on other types of activity. The response to amyl acetate varied between experiments. Blocking the nostrils with wax abolished the response. Some implications of these results are discussed briefly.