Auditory-olfactory integration: congruent or pleasant sounds amplify odor pleasantness

Even though we often perceive odors while hearing auditory stimuli, surprisingly little is known about auditory-olfactory integration. This study aimed to investigate the influence of auditory cues on ratings of odor intensity and/or pleasantness, with a focus on 2 factors: "congruency" (Experiment 1) and the "halo/horns effect" of auditory pleasantness (Experiment 2). First, in Experiment 1, participants were presented with congruent, incongruent, or neutral sounds before and during the presentation of odor. Participants rated the odors as being more pleasant while listening to a congruent sound than while listening to an incongruent sound. In Experiment 2, participants received pleasant or unpleasant sounds before and during the presentation of either a pleasant or unpleasant odor. The hedonic valence of the sounds transferred to the odors, irrespective of the hedonic tone of the odor itself. The more the participants liked the preceding sound, the more pleasant the subsequent odor became. In contrast, the ratings of odor intensity appeared to be little or not at all influenced by the congruency or hedonic valence of the auditory cue. In conclusion, the present study for the first time provides an empirical demonstration that auditory cues can modulate odor pleasantness.     

Facial EMG responses to odors in solitude and with an audience

Two experiments were undertaken to examine whether facial responsesto odors correlate with the hedonic odor evaluation. Experiment1 examined whether subjects (n = 20) spontaneously generatedfacial movements associated with odor evaluation when they aretested in private. To measure facial responses, EMG was recordedover six muscle regions (M. corrugator supercilii, M. procerus,M. nasalis, M. levator, M. orbicularis oculi and M. zygomaticusmajor) using surface electrodes. In experiment 2 the experimentalgroup (n = 10) smelled the odors while they were visually inspectedby the experimenter sitting in front of the test subjects. Thecontrol group (n = 10) performed the same experimental conditionas those subjects participating in experiment 1. Facial EMGover four mimetic muscle regions (M. nasalis, M. levator, M.zygomaticus major, M. orbicularis oculi) was measured whilesubjects smelled different odors. The main findings of thisstudy may be summarized as follows: (i) there was no correlationbetween valence rating and facial EMG responses; (ii) pleasantodors did Dot evoke smiles when subjects smelled the odors inprivate; (iii) in solitude, highly concentrated malodors evokedfacial EMG reactions of those mimetic muscles which are mainlyinvolved in generating a facial display of disgust; (iv) thosesubjects confronted with an audience showed stronger facialreactions over the periocular and cheek region (indicative ofa smile) during the smelling of pleasant odors than those wbosmelled these odors in private; (v) those subjects confrontedwith an audience showed stronger facial reactions over the M.nasalis region (indicative of a display of disgust) during thesmelling of malodors than those who smelled the malodors inprivate. These results were taken as evidence for a more socialcommunicative function of facial displays and strongly mitigatesthe reflexive-hedonic interpretation of facial displays to odorsas supposed by Steiner.     

Odors: implicit memory and performance effects.

In order to assess the influence of odors on human performance and implicit memory for odors, 108 subjects completed a variety of tests in weakly scented (jasmine, lavender or odorless) rooms without having been made aware of the odor. After a 30 min interval the subjects were shown slides of different surroundings, including the room they had been in, and were requested to rate how well a set of 12 odors, including a blank, would fit to these surroundings. Half of these contexts contained visual cues related to two of the presented odors (leather and coffee). After the rating of fit the subjects had to rate the odors for pleasantness, were asked to identify the odors with their correct names and to tell where and when they had last smelled these odors. One subject remembered smelling the odor (jasmine) in the room and was discarded from the analysis of the results for the rating of fit. None of the others reported recollection of the experimental odors. The results showed that in general jasmine had a negative and lavender a positive effect on test performance. If an odor-related visual cue was present in the context, the related odor was always rated highest in fit to that context. Furthermore, the subjects working in rooms with an odor subsequently assigned this odor to the visual context of that room to a significantly higher degree than subjects working in rooms with different odors. Since none of the subjects reported that they had smelled the odor in the rooms where performance testing took place, it was concluded that the memory for these odors was implicit. Further analysis showed that such memory was only found in subjects who were unable to supply the right name for the odor. The possible consequences of this latter finding for understanding the relationship between sensory (episodic) and semantic odor memory are discussed.     

Measures of human olfactory perception during pregnancy

Although considerable anecdotal evidence suggests that pregnancy affects olfactory sensitivity, scientific evidence is limited and inconclusive. Whereas hedonic ratings are affected by pregnancy, odor identification is not. The aim of the current study was to examine odor perception in women across pregnancy and in the postpartum period. One hundred nonsmoking women who were pregnant, postpartum, or had never been pregnant were tested on the University of Pennsylvania Smell Identification Test (UPSIT). Intensity ratings and scratch patterns were collected as potential indicators of sensitivity, and participants rated the odors' pleasantness. Participants also rated their own sense of smell. Mean UPSIT scores did not differ significantly across groups indicating no difference in odor identification. Trends in planned comparisons suggested that in the first trimester, odors were rated as more intense and less pleasant. In the first trimester, women scratched the odor strips significantly fewer times. Consistent with previous reports, 90% of pregnant women reported that specific odors smelled less pleasant and 60% reported that some odors smelled more pleasant. Although nearly two-thirds of pregnant women rated their olfactory sensitivity to be enhanced during pregnancy and overall pregnant women's self-rated olfactory sensitivity was higher than controls', self-ratings were not correlated with UPSIT scores nor odor intensity ratings. These results suggest that these and previous findings may reflect the fact that the effect of pregnancy on olfaction is small and inconsistent.     

Changes in odor quality discrimination following recovery from olfactory nerve transection

Following recovery from olfactory nerve transection, animals regain their ability to discriminate between odors. Odor discrimination is restored after new neurons establish connections with the olfactory bulb. However, it is not known if the new connections alter odor quality perception. To address this question, 20 adult hamsters were first trained to discriminate between cinnamon and strawberry odors. After reaching criterion (> or = 90% correct response), half of the animals received a bilateral nerve transection (BTX) and half a surgical sham procedure. Animals were not tested again until day 40, a point in recovery when connections are re-established with the bulb. When BTX animals were tested without food reinforcement, they could not perform the odor discrimination task. Sham animals, however, could discriminate, demonstrating that the behavioral response had not been extinguished during the 40 day period. When reinforcement was resumed, BTX animals were able to discriminate between cinnamon and strawberry after four test sessions. In addition, their ability to discriminate between these two familiar odors was no different than that of BTX and sham animals tested with two novel odors, baby powder and coffee. These findings suggest that, after recovery from nerve transection, there are alterations in sensory perception and that restoration of odor quality discrimination requires that the animal must again learn to associate individual odor sensations with a behavioral response.      

Olfactory cues modulate facial attractiveness

We report an experiment designed to investigate whether olfactory cues can influence people's judgments of facial attractiveness. Sixteen female participants judged the attractiveness of a series of male faces presented briefly on a computer monitor using a 9-point visual rating scale. While viewing each face, the participants were simultaneously presented with either clean air or else with 1 of 4 odorants (the odor was varied on a trial-by-trial basis) from a custom-built olfactometer. We included 2 pleasant odors (geranium and a male fragrance) and 2 unpleasant odors (rubber and body odor) as confirmed by pilot testing. The results showed that the participants rated the male faces as being significantly less attractive in the presence of an unpleasant odor than when the faces were presented together with a pleasant odor or with clean air (these conditions did not differ significantly). These results demonstrate the cross-modal influence that unpleasant odors can have on people's judgments of facial attractiveness. Interestingly, this pattern of results was unaffected by whether the odors were body relevant (the body odor and the male fragrance) or not (the rubber and geranium odors).     

The signal function of thematically (In)congruent ambient scents in a retail environment

An odor emitted by an object signals the presence of that object and may draw attention to it. Can odors that are not actually emitted by an object also function as a signal? We investigated whether the degree of thematic congruency between an ambient odor and a magazine affected magazine sales in a retail store. We selected two odors: a grass odor, congruent with soccer, animal/nature and gardening magazines; and a sunflower odor, congruent with personal care and women's magazines. In a field study in three bookstores, the ambient odors did not increase sales for thematically congruent magazines, nor did they decrease sales for incongruent magazines. Several explanations for these unexpected findings are discussed.     

Differential foraging in presence of predator and conspecific odors in bank voles: a field enclosure study

Bank voles detect and discriminate other organisms, e.g., predators versus conspecifics, based on olfactory clues like urine, feces, or scent marks. The present field enclosure experiment was set up to determine whether scented food is sufficient for inducing such differentiated reactions in bank voles. Fourteen different odors were tested, divided into four categories: (1) terrestrial predators, (2) avian predators, (3) conspecifics, and (4) neutral origin (human, dog, solvent-treated, and unscented). The odors were extracted with methanol from pellets or feces, or were an artificial substitute. To test the influence of odors on the foraging behavior of voles, artificial food patches containing scented feeders were used. The feeders were cleaned and refilled daily with a calibrated amount of food sprayed with 1 ml of a different odor randomly chosen. The patches were also equipped with monochromatic cameras to monitor the occurrences of voles around the feeders. Both the visits at scented feeders and food consumption were analyzed for each odor. The results show that relative to their foraging activity on unscented food, bank voles significantly reduce it on food scented with odors of terrestrial predators, and increase it in the presence of conspecific odors. Their foraging activity was not affected by neutral scent, nor by scents from avian predators. These results prove that bank voles react to scented food in a way that varies according to the source of odor, even in the semi-natural conditions of an outdoor enclosure.     

Gonadal control of male scent in the tarmarin Saguinus fuscicollis (Callitrichidae, Primates)

Spontaneous responses to the scent marks of male Saguinus fuscicolliswere tested in experiments where the subjects received a choicebetween scent of intact males and castrated males. Male tamarinsdiscriminated between the odors of intact and castrated males,both on the basis of fresh individual scent marks and on thebasis of marks from several donors pooled in methanol. Theirdiscriminatory response to pooled odors was more pronouncedthan that to individual odors. Females discriminated betweenintact and castrated male scents only on the basis of pooledsamples. This result shows that castration causes changes inthe odor of males. The difference in the response of the tamarinsto individual scent marks as compared to pooled samples is discussedin terms of quantitative changes caused by extraction and ofindividual recognition.     

Judgement of odor intensity is influenced by subjects' knowledge of the odor source

Odor perception, including intensity, is affected by knowledge of odor source. For 76 subjects tested with 24 everyday odorants, ratings of intensity, pleasantness and familiarity were enhanced when subjects either could identify the odor source themselves or were provided with the name by the experimenter. Ratings were highest when subjects judged that the names provided matched their own perception, suggesting an interaction between individuals' cognitive representation of odors and their immediate perceptual experience.     

Scent marking by voles in response to predation risk: a field-laboratory validation

Predators use scent to locate their prey, and prey animals oftenalter their behavior in response to predation risk. I testedthe hypothesis that voles would decrease their frequency ofscent marking in response to predation risk. I conducted trialsin which prairie voles, Microtus ochrogaster, and woodland voles,M. pinetorum, were allowed to scent mark ceramic tiles placedin their runways in the field. The tiles were subjected to oneof three treatments: scented with odor from mink, Mustela vison(a rodent predator); rabbit, Oryctolagus cuniculus (a nonpredatormammal control); and no odor (control). No significant differenceswere found in the frequency of scent marking in response tothe three treatments for either species. To validate that volesdid not decrease their scent marking in response to predationrisk, I brought male prairie voles from the field site intothe laboratory and allowed them to scent mark white paper substratetreated with mink odor, rabbit odor, or no odor. No significantdifferences were found in the frequency of scent marks in responseto the three treatments. These results differ from what waspredicted based on laboratory studies with other species ofrodents that show avoidance, reproductive suppression, decreasedactivity, and reduced scent marking in response to odors ofpredators. Voles appear to scent mark different substrates andunder a wide variety of social and environmental situations,and this is not influenced by the presence of odor from a predator.     

Odor-taste interactions: effects of attentional strategies during exposure

Through repeated pairings with a tastant such as sucrose, odors are able to take on the tastant's qualities, e.g. by becoming more sweet smelling. When such odors are subsequently experienced with a sweet tastant in solution, the mixture is often given a higher sweetness rating than the tastant alone. Odor-induced taste enhancement appears to be sensitive to whether an odor-taste combination is viewed analytically as a set of discrete qualities, or synthetically as a flavor. The present research attempted to determine if adoption of these different perceptual approaches during co-exposure with sucrose would influence the extent to which an odor would become sweet smelling and subsequently enhance sweetness intensity. In Experiment 1, subjects received multiple exposures to mixtures of sucrose with low sweetness, low familiarity odors or, as a control, the odors and sucrose solutions separately. Two groups that received mixtures made intensity ratings that promoted either synthesis or analysis of the individual elements in the mixtures. The odors became sweeter smelling irrespective of group. Only adopting a synthetic strategy produced odors that enhanced sweetness in solution. However, these effects were also shown with a 'non-exposed' control odor. This could be accounted for if the single co-exposure with sucrose that all odors received in the pre-test was able to produce sweeter odors. A second experiment confirmed this prediction. Thus, while even a single co-exposure with sucrose is sufficient to produce a sweeter odor, the adoption of a synthetic perceptual strategy during the co-exposure is necessary to produce an odor that will enhance sweetness. These data are consistent with associative leaning accounts of how odors take on taste qualities and also support the interpretation that these effects reflect the central integration of odors and tastes into flavors.     

Attentional modulation of central odor processing

Two studies were conducted to investigate the influence of attention on the components of the chemosensory event-related potential (CSERP). In the first study the odors linalool and eugenol were delivered to six male subjects, in the second study three male and two female subjects were presented with their own body odor (axillary hair) and the body odor of a same sex donor. In both studies the odors were presented in an oddball paradigm under ignore and attend conditions via a constant- flow olfactometer. In the ignore condition attention was diverted from the odors with a distractor task, while in the attend condition the subjects were asked to respond to the infrequently occurring odor. In both studies the allocation of attention led to a decrease in the latency of the early components (N1, P2, N2) and to an increase in the amplitude of the late positivities. The modulation of the early components suggests that attentional gating in olfaction might already be effective at an early processing level.      

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.     

Effects of context on judgements of odor intensities in humans

This study evaluated whether the intensity of previously smelled odors could unintentionally influence the subsequent judgement of odor intensity. The predicted context effect was based on the adaptation- level theory. Before and 25 min after either WEAK or STRONG biasing odor concentrations, 51 subjects were required to rate the intensity of 10 different odor concentrations of California Orange Oil. After the WEAK bias, subjects judged the odor intensity as being stronger than they did after the STRONG bias. Thus the intensity of odors smelled 25 min earlier can unintentionally influence subsequent odor intensity judgement. The findings are discussed in the light of two alternative explanations, namely, a central implicit memory process and a stimulus- level-based change at the peripheral level.      

Ambient odor's effect on creativity, mood, and perceived health

In two sessions held 1 week apart, subjects completed a performancetask involving creativity, four personality tests, and questionnairesconcerning their mood, perceived health, and perceptions ofthe testing environment. In one session the testing room wasscented with lemon, lavender or dimethyl sulfide (DMS); in theother session it was unscented. There were 15 women and 15 menin each odor conditon. Fewer health symptoms were reported inthe lemon conditon on scented compared to unscented days. Subjectsin the DMS group were in a less pleasant mood than those inthe lavender group on both scented and unscented days; the orderin which subjects were exposed to DMS played a role in the moodfindings. There were no significant differences in feelingsof arousal or control. The room was rated as smelling less pleasanton scented compared to unscented days by subjects in the DMScondition. Differences in creativity performance were not significant,but relationships emerged between personality traits and theeffect of odor on task performance. These findings are discussedin terms of how associations and expectations concerning odorsmay play a role in odor's effect on humans.     

Body odor evoked potentials: a new method to study the chemosensory perception of self and non‐self in humans

A new method will be presented which allows the perception of body odors in humans to be studied objectively. The analysis of body odor‐evoked potentials was used to investigate if and how the human brain is able to differentiate self from non‐self body odor for the first time. Six subjects (three females) participated in two experimental sessions. In each session, two body odors (axillary hair) were presented within an olfactory oddball paradigm. One of the odors was collected from the subject and the other from an odor donor of the same sex. In the first session the subjects' attention was distracted to a secondary task (passive paradigm), in the second session the subjects were asked to actively differentiate the odors (active paradigm). For the EEG recordings the odors were presented within a constantly flowing airstream. The results show that the subjects could hardly differentiate the body odors subjectively. However, it could be demonstrated that the central nervous processing of one's own odor was faster than the processing of the chemosensory non‐self signal. Moreover, in the active paradigm, the potentials appeared to be larger when the subjects perceived their own body odor. The conclusion is reached that the measurement of chemosensory event‐related potentials (CSERP) is the method of choice for the investigation of HLA‐associated body odors. This revised version was published online in July 2006 with corrections to the Cover Date.     

The Emotional Distinctiveness of Odor-evoked Memories

A modified paired-associate learning paradigm was used to testwhether odors or verbal odor labels evoked more emotional memories.Subjects were presented with emotionally positive and negativepaintings (to-be-remembered items) in association with positiveand negative odors and odor labels. Painting recall and associatedemotional experience were tested after 48 h. Odor-evoked memorieswere found to be more emotional than verbally cued memorieson a variety of measures. Moreover, if the cue for recall (odoror label) was hedonically congruent with the painting to beremembered, memory for original emotional experiences was enhanced.The findings are discussed within a general cognitive frameworkand implications for using odors to dissociate the emotionaland representational aspects of memory are addressed. Chem.Senses 20: 517–528, 1995.     

Cross-Cultural Color-Odor Associations

Colors and odors are associated; for instance, people typically match the smell of strawberries to the color pink or red. These associations are forms of crossmodal correspondences. Recently, there has been discussion about the extent to which these correspondences arise for structural reasons (i.e., an inherent mapping between color and odor), statistical reasons (i.e., covariance in experience), and/or semantically-mediated reasons (i.e., stemming from language). The present study probed this question by testing color-odor correspondences in 6 different cultural groups (Dutch, Netherlands-residing-Chinese, German, Malay, Malaysian-Chinese, and US residents), using the same set of 14 odors and asking participants to make congruent and incongruent color choices for each odor. We found consistent patterns in color choices for each odor within each culture, showing that participants were making non-random color-odor matches. We used representational dissimilarity analysis to probe for variations in the patterns of color-odor associations across cultures; we found that US and German participants had the most similar patterns of associations, followed by German and Malay participants. The largest group differences were between Malay and Netherlands-resident Chinese participants and between Dutch and Malaysian-Chinese participants. We conclude that culture plays a role in color-odor crossmodal associations, which likely arise, at least in part, through experience.     

The bed nucleus of the stria terminalis is critical for sexual solicitation, but not for opposite-sex odor preference, in female Syrian hamsters

Successful reproduction in vertebrates depends critically upon a suite of precopulatory behaviors that occur prior to mating. In Syrian hamsters (Mesocricetus auratus), these behaviors include vaginal scent marking and preferential investigation of male odors. The neural regulation of vaginal marking and opposite-sex odor preference likely involves an interconnected set of steroid-sensitive nuclei that includes the medial amygdala (MA), the bed nucleus of the stria terminalis (BNST), and the medial preoptic area (MPOA). For example, lesions of MA eliminate opposite-sex odor preference and reduce overall levels of vaginal marking, whereas lesions of MPOA decrease vaginal marking in response to male odors. Although BNST is densely interconnected with both MA and MPOA, little is known about the role of BNST in female precopulatory behaviors. To address this question, females received either bilateral, excitotoxic lesions of BNST (BNST-X) or sham lesions (SHAM), and were tested for scent marking and for investigatory responses to male and female odors. Whereas SHAM females vaginal marked more to male odors than female odors on two days of the estrous cycle, BNST-X females marked at equivalent levels to both odors. This deficit is not due to alterations in social odor investigation, as both BNST-X and SHAM females investigated male odors more than female odors. Finally, BNST lesions did not generally disrupt the cyclic changes in reproductive behaviors that occur across the estrous cycle. Taken together, these results demonstrate that BNST is critical for the normal expression of solicitational behaviors by females in response to male odor stimuli.