Allosteric modulation of a human odorant receptor.

Odor perception is first determined by how the myriad of environmental volatiles are detected at the periphery of the olfactory system. The combinatorial activation of dedicated odorant receptors generates enough encoding power for the discrimination of tens of thousands of odorants. Recent studies have revealed that odorant receptors undergo widespread inhibitory modulation of their activity when presented with mixtures of odorants, a property likely required to maintain discrimination and ensure sparsity of the code for complex mixtures. Here, we establish the role of human OR5AN1 in the detection of musks and identify distinct odorants capable of enhancing its activity in binary mixtures. Chemical and pharmacological characterization indicate that specific α-ß unsaturated aliphatic aldehydes act as positive allosteric modulators. Sensory experiments show decreased odor detection threshold in humans, suggesting that allosteric modulation of odorant receptors is perceptually relevant and likely adds another layer of complexity to how odors are encoded in the peripheral olfactory system.

Exogenous capture of visual spatial attention by olfactory-trigeminal stimuli.

The extent to which a nasal whiff of scent can exogenously orient visual spatial attention remains poorly understood in humans. In a series of seven studies, we investigated the existence of an exogenous capture of visual spatial attention by purely trigeminal (i.e., CO2) and both olfactory and trigeminal stimuli (i.e., eucalyptol). We chose these stimuli because they activate the trigeminal system which can be considered as an alert system and are thus supposedly relevant for the individual, and thus prone to capture attention. We used them as lateralized cues in a variant of a visual spatial cueing paradigm. In valid trials, trigeminal cues and visual targets were presented on the same side whereas in invalid trials they were presented on opposite sides. To characterize the dynamics of the cross-modal attentional capture, we manipulated the interval between the onset of the trigeminal cues and the visual targets (from 580 to 1870 ms). Reaction times in trigeminal valid trials were shorter than all other trials, but only when this interval was around 680 or 1170 ms for CO2 and around 610 ms for eucalyptol. This result reflects that both pure trigeminal and olfactory-trigeminal stimuli can exogenously capture humans' spatial visual attention. We discuss the importance of considering the dynamics of this cross-modal attentional capture.

Factors affecting flavor perception in space: Does the spacecraft environment influence food intake by astronauts?

The intention to send a crewed mission to Mars involves a huge amount of planning to ensure a safe and successful mission. Providing adequate amounts of food for the crew is a major task, but 20 years of feeding astronauts on the International Space Station (ISS) have resulted in a good knowledge base. A crucial observation from the ISS is that astronauts typically consume only 80% of their daily calorie requirements when in space. This is despite daily exercise regimes that keep energy usage at very similar levels to those found on Earth. This calorie deficit seems to have little effect on astronauts who spend up to 12 months on the ISS, but given that a mission to Mars would take 30 to 36 months to complete, there is concern that a calorie deficit over this period may lead to adverse effects in crew members. The key question is why astronauts undereat when they have a supply of food designed to fully deliver their nutritional needs. This review focuses on evidence from astronauts that foods taste different in space, compared to on Earth. The underlying hypothesis is that conditions in space may change the perceived flavor of the food, and this flavor change may, in turn, lead to underconsumption by astronauts. The key areas investigated in this review for their potential impact on food intake are the effects of food shelf life, physiological changes, noise, air and water quality on the perception of food flavor, as well as the link between food flavor and food intake.

Swiss identity smells like chocolate: Social identity shapes olfactory judgments.

There is extensive evidence that social identities can shape people's attitudes and behavior, but what about sensory judgments? We examined the possibility that social identity concerns may also shape the judgment of non-social properties-namely, olfactory judgment. In two experiments, we presented Swiss and non-Swiss participants with the odor of chocolate, for which Switzerland is world-famous, and a control odor (popcorn). Swiss participants primed with Swiss identity reported the odor of chocolate (but not popcorn) as more intense than non-Swiss participants (Experiments 1 and 2) and than Swiss participants primed with individual identity or not primed (Experiment 2). The self-reported intensity of chocolate smell tended to increase as identity accessibility increased-but only among Swiss participants (Experiment 1). These results suggest that identity priming can counter-act classic sensory habituation effects, allowing identity-relevant smells to maintain their intensity after repeated presentations. This suggests that social identity dynamically influences sensory judgment. We discuss the potential implications for models of social identity and chemosensory perception.

A hedonically complex odor mixture produces an attentional capture effect in the brain.

A counter-intuitive property of many pleasant and attractive stimuli is that they are hedonically complex, containing both pleasant and unpleasant components. A striking example is the floral scent of natural jasmine, which may contain more than 6% of indole, a pure chemical which is usually rated as unpleasant. Using fMRI we investigate the hypothesis that the interaction between the pleasant and unpleasant components in the hedonically complex natural jasmine produces an attentional capture effect in the brain. First, to localize brain areas involved in selective attention to odor, we compared neural activity in response to jasmine without indole when participants explicitly and selectively attended to either its pleasantness or intensity, with neural activity when no selective attention was required. We then show that the superior frontal gyrus has increased activity both when selective attention is being paid to jasmine without indole, and also when no selective attention is required but an unpleasant component is added to it to produce a hedonically complex mixture. The attentional capture effect in the superior frontal gyrus by the mixture was related to the hedonic complexity of the mixture across subjects; could not be explained by salience, intensity, or pleasantness; and was specific to the superior frontal gyrus in that it was not found in other prefrontal areas activated by selective attention. The investigation supports the new hypothesis that the affective potency of stimuli with mixed pleasant and unpleasant components is related at least in part to the recruitment of mechanisms in the brain involved in attentional capture and enhancement.

Sequential unfolding of novelty and pleasantness appraisals of odors: evidence from facial electromyography and autonomic reactions.

We investigated the effects of odors on appraisal processes and consequent emotional responses. The main goal was to test whether an odor is detected as novel or familiar before it is evaluated as pleasant or unpleasant. Participants performed a recognition task in which they were presented with pairs of unpleasant or pleasant odors (sample and target odors). Within a pair, the sample and target were either identical or different to assess participants' novelty detection; unpleasant and pleasant target odors were contrasted to examine participants' appraisal of intrinsic pleasantness. We measured facial expressions using electromyography and physiological reactions using electrocardiogram and electrodermal activity in response to odors. The earliest effects on facial muscles and heart rate occurred in response to novelty detection. Later effects on facial muscles and heart rate were related to pleasantness evaluation. This study is the first to demonstrate the existence of a sequence of appraisal checks for odors eliciting emotional reaction.

Selective attention to affective value alters how the brain processes olfactory stimuli.

How does selective attention to affect influence sensory processing? In a functional magnetic resonance imaging investigation, when subjects were instructed to remember and rate the pleasantness of a jasmine odor, activations were greater in the medial orbito-frontal and pregenual cingulate cortex than when subjects were instructed to remember and rate the intensity of the odor. When the subjects were instructed to remember and rate the intensity, activations were greater in the inferior frontal gyrus. These top-down effects occurred not only during odor delivery but started in a preparation period after the instruction before odor delivery, and continued after termination of the odor in a short-term memory period. Thus, depending on the context in which odors are presented and whether affect is relevant, the brain prepares itself, responds to, and remembers an odor differently. These findings show that when attention is paid to affective value, the brain systems engaged to prepare for, represent, and remember a sensory stimulus are different from those engaged when attention is directed to the physical properties of a stimulus such as its intensity. This differential biasing of brain regions engaged in processing a sensory stimulus depending on whether the cognitive demand is for affect-related versus more sensory-related processing may be an important aspect of cognition and attention. This has many implications for understanding the effects not only of olfactory but also of other sensory stimuli.

How pleasant and unpleasant stimuli combine in different brain regions: odor mixtures.

Many affective stimuli are hedonically complex mixtures containing both pleasant and unpleasant components. To investigate whether the brain represents the overall affective value of such complex stimuli, or the affective value of the different components simultaneously, we used functional magnetic resonance imaging to measure brain activations to a pleasant odor (jasmine), an unpleasant odor (indole), and a mixture of the two that was pleasant. In brain regions that represented the pleasantness of the odors such as the medial orbitofrontal cortex (as shown by activations that correlated with the pleasantness ratings), the mixture produced activations of similar magnitude to the pleasant jasmine, but very different from the unpleasant indole. These regions thus emphasize the pleasant aspects of the mixture. In contrast, in regions representing the unpleasantness of odors such as the dorsal anterior cingulate and midorbitofrontal cortex the mixture produced activations that were relatively further from the pleasant component jasmine and closer to the indole. These regions thus emphasize the unpleasant aspects of the mixture. Thus mixtures that are found pleasant can have components that are separately pleasant and unpleasant, and the brain can separately and simultaneously represent the positive and negative hedonic value of a complex affective stimulus that contains both pleasant and unpleasant olfactory components. This type of representation may be important for affective decision making in the brain in that separate representations of different affective components of the same sensory stimulus may provide the inputs for making a decision about whether to choose the stimulus or not.

Cognitive modulation of olfactory processing.

We showed how cognitive, semantic information modulates olfactory representations in the brain by providing a visual word descriptor, "cheddar cheese" or "body odor," during the delivery of a test odor (isovaleric acid with cheddar cheese flavor) and also during the delivery of clean air. Clean air labeled "air" was used as a control. Subjects rated the affective value of the test odor as significantly more unpleasant when labeled "body odor" than when labeled "cheddar cheese." In an event-related fMRI design, we showed that the rostral anterior cingulate cortex (ACC)/medial orbitofrontal cortex (OFC) was significantly more activated by the test stimulus and by clean air when labeled "cheddar cheese" than when labeled "body odor," and the activations were correlated with the pleasantness ratings. This cognitive modulation was also found for the test odor (but not for the clean air) in the amygdala bilaterally.