Differential neural responses evoked by orthonasal versus retronasal odorant perception in humans

Odors perceived through the mouth (retronasally) as flavor are referred to the oral cavity, whereas odors perceived through the nose (orthonasally) are referred to the external world. We delivered vaporized odorants via the orthonasal and retronasal routes and measured brain response with fMRI. Comparison of retronasal versus orthonasal delivery produced preferential activity in the mouth area at the base of the central sulcus, possibly reflecting olfactory referral to the mouth, associated with retronasal olfaction. Routes of delivery produced differential activation in the insula/operculum, thalamus, hippocampus, amygdala, and caudolateral orbitofrontal cortex in orthonasal > retronasal and in the perigenual cingulate and medial orbitofrontal cortex in retronasal > orthonasal in response to chocolate, but not lavender, butanol, or farnesol, so that an interaction of route and odorant may be inferred. These findings demonstrate differential neural recruitment depending upon the route of odorant administration and suggest that its effect is influenced by whether an odorant represents a food.     

Identification of air phase retronasal and orthonasal odorant pairs

Identifications (IDs) of paired retronasal and orthonasal odorants were studied, with stimuli limited to air phase. Odorants were liquid extracts of plant materials, sold as food flavorings, matched by each subject both for retronasal-only and orthonasal-only air phase intensities and then learned to 100% correct veridical name retronasal-only and orthonasal-only IDs. Subjects were tested for ID of (a) retronasal-only and orthonasal-only odorants, (b) homogeneously paired odorant (the same odorant in retronasal and orthonasal locations), and (c) heterogeneously paired odorants (different odorants in retronasal and orthonasal locations). Paired odorants were presented in two different sequences: retronasal location odorant smelled first or orthonasal location odorant smelled first. IDs were reported after odorants were removed. Results were as follows: (a) no significant differences between correct ID of odorants when in retronasal-only versus orthonasal-only locations, although percent correct IDs were lower for half the retronasal-only location odorants; (b) correct ID of a homogeneously paired odorant equaled or exceeded its unpaired ID, with two successive, identical IDs reported on the majority of its trials; (c) with heterogeneous pairs, for all odorants when in the orthonasal location of a pair, correct ID occurred less often than when these odorants were presented orthonasal-only, but for odorants in the retronasal location, correct ID equaled or exceeded retronasal-only correct ID; and (d) perceived order of presentation of heterogeneous pairs was the reverse of the physically presented sequence for both retronasal-first and orthonasal-first conditions. The heterogeneous odorant ID outcome supports the concept that processing of retronasal and orthonasal odorants differ, and the perceived reversal of the presented sequence is in agreement with the importance of recency in odorant memory.     

Comparison of the orthonasal and retronasal detection thresholds for carbon dioxide in humans

Several studies have investigated the orthonasal detection threshold for carbon dioxide (CO(2)) in humans. The aim of current study was to investigate whether 24 healthy young subjects exhibited differences of CO(2) detection thresholds during orthonasal or retronasal stimulation. As nasal mucosa is believed to desensitize to CO(2) concentrations at or below 4% (v/v) during expiration, the second aim of the study was to explore the influence during nasal versus oral breathing on the detection thresholds. CO(2) stimuli of varying concentrations and a duration of 1000 ms were applied with an air-dilution olfactometer in either the anterior nasal cavity or the nasopharynx during nasal respectively oral breathing. In these 4 conditions, the mean CO(2) detection thresholds using the staircase forced-choice procedure were between 3.9% and 5.3% (v/v). Statistical analysis revealed a significant difference between orthonasal and retronasal stimulation. The CO(2) detection threshold was lower in retronasal stimulation. The nasopharyngeal mucosa is more sensitive to perithreshold CO(2) stimuli than the nasal mucosa. The breathing route had no influence on the detection thresholds. The results of this study indicate that the natural contact of the nasal mucosa with approximately 4% (v/v) CO(2) during nasal expiration does not influence CO(2) detection thresholds.     

Mechanical obstruction of the olfactory cleft reveals differences between orthonasal and retronasal olfactory functions

Following up on recent observations in patients with nasal polyposis (NP), the present study aimed to investigate whether a mechanical obstruction of the anterior olfactory cleft (OC) would produce differential effects on orthonasal and retronasal olfactory functions. To this end, we studied 33 healthy subjects in a randomized trial. Sponges with high content of saline were either placed in the OC or on the respiratory epithelium, such that this was blinded to both subject and observer. The results indicated that orthonasal (P = 0.04) but not retronasal (P = 0.15) olfactory identification ability was lower when the OC was blocked. This confirms the idea that differences between orthonasal and retronasal olfactory functions, as observed in NP patients, are, at least to some degree, due to mechanical obstruction of the anterior portion of the OC. The present data also suggest that mechanical obstruction is a means to induce reversible hyposmia void of side effects which can be performed in a blinded fashion. This might become a valuable model of hyposmia for future investigations.     

The role of congruency in retronasal odor referral to the mouth

Referral of retronasal odors to the mouth is a fundamental phenomenon of flavor perception. A previous study from this laboratory provided evidence that, contrary to prior speculation, taste rather than touch was the primary factor in retronasal odor referral. The present study further investigated this question by studying the role of congruency between taste and odor on retronasal odor referral under conditions that mimicked natural food consumption. Subjects performed odor localization tasks after sampling gelatin stimuli that contained various congruent and incongruent tastes-odor combinations. The results showed that when a congruent taste was added, referral to the oral cavity and tongue were significantly enhanced. In addition, the data also indicate that the degree of congruency between taste and odor may modulate the degree of odor referral to the mouth. These findings suggest that odor referral is maximized when congruent flavor dimensions are combined to trigger perceptual "flavor objects" that represent known or potential foods. The results are discussed in terms of the factors that play a role in the retronasal odor referral as well as the potential neural mechanisms that may underlie it.     

The relationship between the side tastes of masking stimuli and masking in binary mixtures

Subjects estimated the sweetness of HOH, NaCl, Q₂SO₄, sucroseand of the binary mixture of the three compounds. Another groupof subjects made independent estimates of the bitterness andsaltiness of HOH, Q₂SO₄, NaCl and of the binary mixtures ofNaCl and Q₂SO₄. Apart from their specific main taste, the stimuliappear to also possess a side taste in the modality equal tothe suppressed component, with the exception of Q₂SO₄ whichshowed no sweet side taste. The side tastes are negatively relatedto masking, thus the higher a specific side taste, the lowerthe amount of suppression of that taste in the binary mixture.Apparently, the subjects add these side tastes of the maskingagents to the specific quality of the suppressed sensationsin the mixture. In this way spurious results of low maskingor even enhancement are produced. The obtained results werecompared to data from a study in the literature on taste masking.     

Axonal effects of capsaicin: an electrophysiological study

The effects of axonally applied capsaicin on the discharge activity and compound action potential of the cat vagus, saphenous and phrenic nerves and the cervical sympathetic trunk were studied under in vivo and in vitro conditions. Application of capsaicin (10(-4) M) to the vagus, saphenous and phrenic nerves resulted in the appearance of intense discharge activity which reached its maximum after 3-4 min and lasted for 15-20 min. Parallel with the increase in discharge activity, elicited by orthodromic activation induced by capsaicin, the amplitudes of the A delta and C components of an antidromically evoked compound action potential were significantly reduced. After the excitatory action of capsaicin vanished, an increase in the latency and duration and a decrease in the amplitudes of the components of the compound action potential were observed which might have led to the development of a local block of impulse propagation. These changes proved to be reversible after the removal of capsaicin from the nerve. Compound action potentials recorded from the saphenous or vagus nerves pretreated with capsaicin 3-5 days before the experiments failed to show any significant changes. It is concluded that upon direct axonal application capsaicin results in the excitation of both A delta and C fibres which is followed by a nonspecific but reversible blockade of impulse propagation. The possible significance of these transient effects of axonally applied capsaicin in term of the development of the highly specific functional impairment occurring a few days after perineural capsaicin treatment remains to be elucidated.     

Context-dependent changes in the perception of odor quality

Ambiguous odor compounds, partly citrus-like and partly woodyin odor character, were seen to change in odor quality whenevaluated in the same session as more prototypical odors. Whentested with characteristically citrus odors, the ambiguous compoundsseemed more woody, and when tested with characteristically woodyodors, the ambiguous odorants were higher in citrus character,an example of perceptual contrast. Response frequency biaseswere ruled out as an explanation for this shift by an experimentin which responses other than citrus and woody ratings wereasked of the subjects during the contextual exposure. Simplesensory adaptation was found to be a potential contributor tothe effect, and a sufficient condition to produce similar shiftsin odor quality. However, adaptation was not a necessary conditionto produce the effect. This was seen in reversed pair experimentsin which the contextual odors were presented after the ambiguousstimuli. The contextual shift was robust—it was obtainedwith different ambiguous odors, contextual (conditioning) odors,numbers of contextual odors, orders of presentation of contextualodors relative to ambiguous odors, scale types, and rating tasksduring the presentation of contextual odors.     

The perception of odor intensity: physics or psychophysics?

The molecular-kinetic theory as applied to the interaction ofgaseous molecules with a solid surface leads to formulae whichare consistent with the results of psychophysical measurementsof perceived odor intensity.     

Dynamical vs. judgmental comparison: hysteresis effects in motion perception

Perceptual comparison was investigated by gradually varying the relative length of two apparent motion paths, and independently determining when an initial percept was lost during the course of attribute change and when an alternative percept emerged. Dynamical comparison was indicated by a range of attribute values for which perception was bistable. Within this range, a percept that lost stability was immediately replaced by an alternative percept. Judgmental comparison was indicated by a range of attribute values for which perception was uncertain. When an initial percept was lost, an alternative percept did not immediately emerge because the alternatives being compared could not be distinguished. Differences in the effects of random noise on dynamical vs. judgmental comparison were demonstrated with computational simulations, and implications are discussed for motion energy models and solutions to the motion correspondence problem.     

Memory psychophysics for chemosensation: perceptual and mental mixtures of odor and taste

Subjects made quantitative judgements of the overall intensityof odor-taste mixtures, which were either presented physically(perceptual estimation) or represented symbolically by pairsof colors that referred to their unmixed constituents (memory-basedestimation). In the latter condition, the mixtures were constructedsubjectively by mixing mentally the remembered representationsof the odor and taste components. Despite the great phenomenaldifference between the two conditions, the pattern of resultswas largely the same for perceived and for mental mixtures.The rule of odor-taste integration was approximately additivein both cases. The invariance of this rule is particularly impressivegiven that all of the stimuli were presented intraorally, sothat the odorants were mislocalized at the mouth. The findingsimply an important role for cognition in chemosensation.     

Performance of mice in an automated olfactometer: odor detection, discrimination and odor memory

Mice were trained on a variety of odor detection and discrimination tasks in 100- or 200-trial sessions using a go, no-go discrete trials operant conditioning procedure. Odors, presented for 1 s on each trial, were generated by an air dilution olfactometer (for threshold tests) and an easily constructed eight-channel liquid dilution unit (for two- and multiple-odor discrimination tasks). Mice rapidly acquired the operant task and demonstrated excellent stimulus control by odor vapors. Their absolute detection threshold for ethyl acetate was similar to that obtained with rats using similar methods. They readily acquired four separate two-odor discrimination tasks and continued to perform well when all eight odors were presented in random order in the same session and when reinforcement probability for correct responding was decreased from 1 to 0.5. Memory for these eight odors, assessed under extinction after a 32 day rest period, was essentially perfect. Time spent sampling the odor on S+ and S- trials was highly correlated with response accuracy. When accuracy was at chance levels (e.g. initial trials on a novel task), stimulus sampling time on both S+ and S- trials was approximately 0.5-0.7 s. As response accuracy increased, sampling time on S+ trials tended to increase and remain higher than sampling time on S- trials.     

New evidence that the MHC influences odor perception in humans: a study with 58 Southern Brazilian students

Increasing evidence suggests a correlation between mate choice, odor preference, and genetic similarity at the Major Histocompatibility Complex (MHC) in a variety of animals, including our species. The MHC is a highly polymorphic group of genes that play an important role in the immunological self/nonself recognition. Its products have been reported to take part on the variety of compounds and reactions that together build an individual's body odor. It has been suggested, therefore, that animals use body odor as a guide to identify possible mates as MHC-similar or MHC-dissimilar from their own genotype. Preference for a MHC-dissimilar partner enhances MHC heterozygosity of an individual's offspring. The possible adaptive advantages are clear: it is a mechanism of avoiding inbreeding and MHC-heterozygous offspring may have enhanced immunocompetence. The aim of this study was to search, in our species, new evidence on the correlation between specificities at HLA-A and HLA-B and assessments of pleasantness regarding specific body odors. HLA is the name for the human MHC. Four olfactory sessions were performed with 58 young Southern Brazilian students, in order to investigate whether assessments of pleasantness of body odors from individuals correlate to a person's HLA phenotype. Body odors were collected via sweat and urine from all participants. Women smelled and scored all male odor samples and men did the same with all female samples. We found a significant correlation only when female smellers evaluated male sweat odors.     

Semantics in action: an electrophysiological study on the use of semantic knowledge for action.

Recent studies have supported close interactions between language and action-related processes, suggesting comparable neural mechanisms. However, relatively little is known about the semantics involved in action planning. The present study investigated the activation of semantic knowledge in meaningful actions by recording event-related potentials (ERPs). Subjects prepared meaningful or meaningless actions with objects and made a semantic categorization response before executing the action. Words presented could be either congruent or incongruent with respect to the goal of the action. Preparation of meaningful actions elicited a larger anterior N400 for words incongruent to the present action goal as compared to congruent words, while no N400 effect was found when subjects prepared meaningless actions. These findings indicate that the preparation of meaningful actions with objects is accompanied by the activation of semantic information representing the usual action goals associated with those objects.     

Investigation of breathing parameters during odor perception and olfactory imagery

Compared with visual and auditory imagery, little is known aboutolfactory imagery. There is evidence that respiration may bealtered by both olfactory perception and olfactory imagery.In order to investigate this relationship, breathing parameters(respiratory minute volume, respiratory amplitude, and breathingrate) in human subjects during olfactory perception and olfactoryimagery were investigated. Fifty-six subjects having normalolfactory function were tested. Nasal respiration was measuredusing a respiratory pressure sensor. Using an experimental blockdesign, we alternately presented odors or asked the subjectsto imagine a given smell. Four different pleasant odors wereused: banana, rose, coffee, and lemon odor. We detected a significantincrease in respiratory minute volume between olfactory perceptionand the baseline condition as well as between olfactory imageryand baseline condition. Additionally we found significant differencesin the respiratory amplitude between imagery and baseline conditionand between odor and imagery condition. Differences in the breathingrate between olfactory perception, olfactory imagery, and baselinewere not statistically significant. We conclude from our resultsthat olfactory perception and olfactory imagery both have effectson the human respiratory profile and that these effects arebased on a common underlying mechanism.     

Decoding visual roughness perception: an fMRI study

Abstract

The neural substrates of tactile roughness perception have been investigated by many neuroimaging studies, while relatively little effort has been devoted to the investigation of neural representations of visually perceived roughness. In this human fMRI study, we looked for neural activity patterns that could be attributed to five different roughness intensity levels when the stimuli were perceived visually, i.e., in absence of any tactile sensation. During functional image acquisition, participants viewed video clips displaying a right index fingertip actively exploring the sandpapers that had been used for the behavioural experiment. A whole brain multivariate pattern analysis found four brain regions in which visual roughness intensities could be decoded: the bilateral posterior parietal cortex (PPC), the primary somatosensory cortex (S1) extending to the primary motor cortex (M1) in the right hemisphere, and the inferior occipital gyrus (IOG). In a follow-up analysis, we tested for correlations between the decoding accuracies and the tactile roughness discriminability obtained from a preceding behavioural experiment. We could not find any correlation between both although, during scanning, participants were asked to recall the tactilely perceived roughness of the sandpapers. We presume that a better paradigm is needed to reveal any potential visuo-tactile convergence. However, the present study identified brain regions that may subserve the discrimination of different intensities of visual roughness. This finding may contribute to elucidate the neural mechanisms related to the visual roughness perception in the human brain.