Individual differences in the chemical senses: is there a common sensitivity?

Taste, smell, and chemical irritation (so-called trigeminal sensation) combine in our daily experience to produce the supramodal sensation of flavor, are processed by partly overlapping neural mechanisms, and show functional interconnectivity in experiments. Given their collaboration in flavor formation and the well-established connections between these senses, it is plausible that polymodal detection mechanisms might contribute to individual differences in measured sensitivity. One would expect the existence of a general chemosensory sensitivity factor to result in associations among taste, smell, and trigeminal stimulation thresholds. Measures of 5 detection thresholds from all the chemical senses were assessed in the same group of young healthy subjects (n=57). An unbiased principal components analysis (PCA) yielded a 2-component solution. Component 1, on which taste thresholds loaded strongly, accounted for 29.4% of the total variance. Component 2, on which the odor and trigeminal lateralization thresholds loaded strongly, accounted for 26.9% of the total variance. A subsequent PCA restricted to a 3-component solution cleanly separated the 3 sensory modalities and accounted for 75% of the total variance. Thus, though there may be a common underlying factor that determines some individual differences in odor and trigeminal lateralization thresholds, a general chemical sensitivity that spans chemosensory modalities seems unlikely.     

Chemosensory cross-modal stroop effects: congruent odors facilitate taste identification

In order to explore the cross-modal cognitive associations between smell and taste, a chemosensory analogue of the Stroop task (Stroop 1935) was developed. Fourteen participants were presented with an odorant and a tastant and asked to identify the tastant as "sweet" or "sour" by pressing 1 of 2 buttons as quickly as possible. Participants were faster to name the taste when it was presented with an odor that was congruent (e.g., strawberry/sweet) than with an incongruent odor (e.g., strawberry/sour). These results support the concept of a high level of cognitive integration between the senses of smell and taste and illustrates occasions of interference between information arising from different sensory systems.     

Modulation of taste sensitivity by GLP-1 signaling

In many sensory systems, stimulus sensitivity is dynamically modulated through mechanisms of peripheral adaptation, efferent input, or hormonal action. In this way, responses to sensory stimuli can be optimized in the context of both the environment and the physiological state of the animal. Although the gustatory system critically influences food preference, food intake and metabolic homeostasis, the mechanisms for modulating taste sensitivity are poorly understood. In this study, we report that glucagon-like peptide-1 (GLP-1) signaling in taste buds modulates taste sensitivity in behaving mice. We find that GLP-1 is produced in two distinct subsets of mammalian taste cells, while the GLP-1 receptor is expressed on adjacent intragemmal afferent nerve fibers. GLP-1 receptor knockout mice show dramatically reduced taste responses to sweeteners in behavioral assays, indicating that GLP-1 signaling normally acts to maintain or enhance sweet taste sensitivity. A modest increase in citric acid taste sensitivity in these knockout mice suggests GLP-1 signaling may modulate sour taste, as well. Together, these findings suggest a novel paracrine mechanism for the regulation of taste function.     

Odor Perception and Beliefs about Risk

Although the perceptual response to environmental odors canbe quite variable, such variation has often been attributedto differences in individual sensitivity. An information-processinganalysis of odor perception, however, treats both the receptionand the subsequent evaluation of odor information as determinantsof the perceptual response. Two experiments investigated whethera factor that influenced the evaluation stage affected the judgementof odor quality and the degree of adaptation to the odor. Peoplewere surveyed in order to measure their tacit perceptions ofthe healthfulness or hazardousness of nine common olfactorystimuli, and the instructional context influenced quality perception.In a second experiment subjects were exposed to an ambient odorunder one of three different conditions, and odorant characterizationinfluenced the degree of adaptation to the odor. Subjects whowere led to believe the odor was a natural, healthy extractshowed adaptation; those told that the odor was potentiallyhazardous showed apparent sensitization; while those told thatthe odor was a common olfactory test odorant showed a mixedpattern: some exhibited adaptation, whereas others showed sensitization.However, detection thresholds obtained before and after exposureshowed adaptation effects that are characteristic of continuousexposure. These findings raise the possibility that cognitivefactors may be modulating the overall sensory perception ofodor exposure (i) for some individuals who exhibit extreme sensitivityto odors and (ii) in situations where adaptation to environmentalodors is expected but does not occur. Chem. Senses 21: 447–458,1996.     

The Gustatory Neural Response Function

With few exceptions, afferent neurons in the various sensory systems respond to wide ranges of stimuli. In those sensory systems for which the stimulus dimensions are understood, the response functions of these neurons may be described; they are usually simple functions with one maximum, although many variations exist. In the chemical senses, the stimulus dimensions are not known, and thus the neural response functions of these neurons have never been described. The present paper presents methods to determine these response functions and the stimulus dimensions for the chemical senses. A tentative response function for taste is developed, and preliminary steps are taken toward disclosing the stimulus dimensions.     

Invertebrate-Inspired sensory-motor systems and autonomous, olfactory-guided exploration

The localization of resources in a natural environment is a multifaceted problem faced by both invertebrate animals and autonomous robots. At a first approximation, locomotion through natural environments must be guided by reliable sensory information. But natural environments can be unpredictable, so from time to time, information from any one sensory modality is likely to become temporarily unreliable. Fortunately, compensating mechanisms ensure that such signals are replaced or disambiguated by information from more reliable modalities. For invertebrates and robots to rely primarily on chemical senses has advantages and pitfalls, and these are discussed. The role of turbulence, which makes tracking a single odor to its source a complex problem, is contrasted with the high-fidelity identification of stimulus quality by the invertebrate chemoreceptor and by artificial sensors.     

TIME-INTENSITY SCALING WITH JUDGES TRAINED TO USE A CALIBRATED SCALE: ADAPTATION, SALTY AND UMAMI TASTES

The Time-Intensity scaling technique was modified for measurements of taste intensity. Instead of using the traditional procedure where intensities were estimated on the judges’ own internalized scales, judges were trained to estimate concentrations directly in millimolar concentration. Such training was seen to increase precision, although the general shapes of the Time-Intensity plots remained the same. The technique was applied to measurement of taste intensity change over time when NaCl or MSG stimuli were held in the mouth. Using this technique, it was possible to measure the relative effects, on taste intensity loss, of stimulus dilution by saliva and sensory adaptation; the latter had the greater effect. Accordingly, the disruption of adaptation was found to be more important than increased salivation when chewing movements were made.     

Multisensory Integration and Behavioral Plasticity in Sharks from Different Ecological Niches

The underwater sensory world and the sensory systems of aquatic animals have become better understood in recent decades, but typically have been studied one sense at a time. A comprehensive analysis of multisensory interactions during complex behavioral tasks has remained a subject of discussion without experimental evidence. We set out to generate a general model of multisensory information extraction by aquatic animals. For our model we chose to analyze the hierarchical, integrative, and sometimes alternate use of various sensory systems during the feeding sequence in three species of sharks that differ in sensory anatomy and behavioral ecology. By blocking senses in different combinations, we show that when some of their normal sensory cues were unavailable, sharks were often still capable of successfully detecting, tracking and capturing prey by switching to alternate sensory modalities. While there were significant species differences, odor was generally the first signal detected, leading to upstream swimming and wake tracking. Closer to the prey, as more sensory cues became available, the preferred sensory modalities varied among species, with vision, hydrodynamic imaging, electroreception, and touch being important for orienting to, striking at, and capturing the prey. Experimental deprivation of senses showed how sharks exploit the many signals that comprise their sensory world, each sense coming into play as they provide more accurate information during the behavioral sequence of hunting. The results may be applicable to aquatic hunting in general and, with appropriate modification, to other types of animal behavior.     

A model of olfactory adaptation and sensitivity enhancement in the olfactory bulb

It has been suggested that the olfactory bulb, the first processing center after the sensory cells in the olfactory pathway, plays a role in olfactory adaptation, odor sensitivity enhancement by motivation and other olfactory psychophysical phenomena. In a mathematical model based on the bulbar anatomy and physiology, the inputs from the higher olfactory centers to the inhibitory cells in the bulb are shown to be able to modulate the response, and thus the sensitivity of the bulb to specific odor inputs. It follows that the bulb can decrease its sensitivity to a pre-existing and detected odor (adaptation) while remaining sensitive to new odors, or increase its sensitivity to interested searching odors. Other olfactory psychophysical phenomena such as cross-adaptation etc. are discussed as well.     

Effects of selective adaptation on coding sugar and salt tastes in mixtures

Little is known about coding of taste mixtures in complex dynamic stimulus environments. A protocol developed for odor stimuli was used to test whether rapid selective adaptation extracted sugar and salt component tastes from mixtures as it did component odors. Seventeen human subjects identified taste components of "salt + sugar" mixtures. In 4 sessions, 16 adapt-test stimulus pairs were presented as atomized, 150-μL "taste puffs" to the tongue tip to simulate odor sniffs. Stimuli were NaCl, sucrose, "NaCl + sucrose," and water. The sugar was 98% identified but the suppressed salt 65% identified in unadapted mixtures of 2 concentrations of NaCl, 0.1 or 0.05 M, and sucrose at 3 times those concentrations, 0.3 or 0.15 M. Rapid selective adaptation decreased identification of sugar and salt preadapted ambient components to 35%, well below the 74% self-adapted level, despite variation in stimulus concentration and adapting time (<5 or >10 s). The 96% identification of sugar and salt extra mixture components was as certain as identification of single compounds. The results revealed that salt-sugar mixture suppression, dependent on relative mixture-component concentration, was mutual. Furthermore, like odors, stronger and recent tastes are emphasized in dynamic experimental conditions replicating natural situations.     

CROSS-MODAL ADDITIVITY OF TASTE AND SMELL

Subjects were simultaneously given subthreshold levels of taste and odor stimuli, delivered orally, for both a commonly paired and an uncommonly paired taste–odor combination. Results indicate cross‐modal summation of subthreshold concentrations of both taste–odor pairs when the olfactory stimulus is delivered orally. Results of control studies suggest that the summation was indeed across modalities, and not due to the taste of the odor compound or the smell of the taste compounds. Furthermore, results indicate that regardless of taste–odor pair commonness, taste and smell can combine in a completely additive fashion (i.e., at threshold detectability when both stimuli are presented simultaneously at 50% threshold level) if the taste–odor pair is presented orally. In several instances, but not all, measured probabilities exceeded those predicted by probability summation, indicating that hyperadditive mixing often occurs, but there do seem to be individual differences. Cross‐modal summation, regardless of taste–odor pair commonness, has broader implications for the development of foods, beverages and pharmaceuticals, especially in masking undesirable tastes and smells.     

Studies on the Chemical Evaluation of Tobacco Quality

Various organoleptic characteristics of cigarette smoke, such as aroma, taste, harshness, offensive odor and taste, strength and dryness, were measured by sensory test using more than fifty kinds of flue-cured tobacco leaves. The results were compared with those of chemical and gas chromatographic analyses of the particulate matter in the smoke. The correlations between each evaluation characteristic and each analytical value were computed and simple equations for tobacco evaluation were set up. The equations obtained were considered to be available for the evaluation of tobacco quality instead of the sensory test, although their applications to lowgrade samples seemed to need further modifications.     

NEUROPHYSIOLOGY AND HUMAN TASTE SENSATIONS

Taste sensations are of primary importance in food flavor. Any attempt to synthesize chemically the flavor of a natural food involves mainly taste active compounds. Many distinct taste sensations can be identified as associated with food compounds. Thirteen different taste sensations are discussed herein. These different taste sensations are differentiated on the basis of stimulus chemistry and peripheral nerve conveying the taste information. Neurophysiological examination of the peripheral nerves involved in taste reveals that the sensory neurons can, in any species, be subdivided into distinct neural groups. These different neural groups respond to distinct classes of chemicals and often display different neurophysiological characteristics. Altogether in four different species, nine functional neural taste groups can be distinguished. In many cases, these neural groups can be taken as analogs for the neural groups assumed to underly human taste sensations. Distinct human taste sensations can be considered to arise from the excitation or inhibition of different neural groups. For certain human taste sensations there are no animal neural analog groups; and for certain neural groups there are no analog human sensations.     

Flavor-Enhanced Modulation of Cerebral Blood Flow during Gum Chewing

Background

Flavor perception, the integration of taste and odor, is a critical factor in eating behavior. It remains unclear how such sensory signals influence the human brain systems that execute the eating behavior.

Methods

We tested cerebral blood flow (CBF) in the frontal lobes bilaterally while subjects chewed three types of gum with different combinations of taste and odor: no taste/no odor gum (C-gum), sweet taste/no odor gum (T-gum), and sweet taste/lemon odor gum (TO-gum). Simultaneous recordings of transcranial Doppler ultrasound (TCD) and near infrared spectrometer (NIRS) were used to measure CBF during gum chewing in 25 healthy volunteers. Bilateral masseter muscle activity was also monitored.

Results

We found that subjects could discriminate the type of gum without prior information. Subjects rated the TO-gum as the most flavorful gum and the C-gum as the least flavorful. Analysis of masseter muscle activity indicated that masticatory motor output during gum chewing was not affected by taste and odor. The TCD/NIRS measurements revealed significantly higher hemodynamic signals when subjects chewed the TO-gum compared to when they chewed the C-gum and T-gum.

Conclusions

These data suggest that taste and odor can influence brain activation during chewing in sensory, cognitive, and motivational processes rather than in motor control.     

A model of the role of adaptation and disadaptation in olfactory receptor neurons: implications for the coding of temporal and intensity patterns in odor signals

Natural odors occur as turbulent plumes resulting in spatiallyand temporally variable odor signals at the chemoreceptor cells.Concentrations can fluctuate widely within discrete packetsof odor and individual packets are very intermittent and unpredictable.Chemoreceptor cells display the temporally dynamic propertiesof adaptation and disadaptation, which serve to alter theirresponses to these fluctuating odor patterns. A computationalmodel, modified from one previously published, was used to investigate,the effect of adaptation and recovery of adaptation (disadaptation)on the spike output of model olfectory receptor cells undernatural stimulus conditions. The response characteristics ofmodel cells were based upon empirically determined dose-response,adaptation, disadaptation and flicker fusion properties of peripheralolfactory cells. The physiological properties of the model cell(adaptation and disadaptation rate and the dose-response relationship)could be modified independently, allowing assessment of therole of each in shaping the responses of the model cell. Completeadaptation and disadaptation time courses ranged from 500 ms(rapid cells) to 10 s (slow cells). The stimuli for the modelcells were quantified odor plume recordings obtained under avariety of biologically relevant flow conditions. As expected,the rapidly adapting model cells displayed different responsecharacteristics than the slowly adapting model cells to identicaltemporal odor profiles. Responses of the model cells dependedupon their adaptation and disadaptation rates, and the frequencycharacteristics of the odor presentation. These results indicatethat adaptation and disadaptation determine the range of concentrationfluctuations over which a particular cell will respond. Thus,these properties function as an olfactory equivalent of a band-passfilter in electronics. This type of filtering has implicationsfor the extraction of information from odor signals, men isthe coding of temporal and intensity features.     

Ca2+-activated K+ currents regulate odor adaptation by modulating spike encoding of olfactory receptor cells

The olfactory system is thought to accomplish odor adaptation through the ciliary transduction machinery in olfactory receptor cells (ORCs). However, ORCs that have lost their cilia can exhibit spike frequency accommodation in which the action potential frequency decreases with time despite a steady depolarizing stimulus. This raises the possibility that somatic ionic channels in ORCs might serve for odor adaptation at the level of spike encoding, because spiking responses in ORCs encode the odor information. Here I investigate the adaptational mechanism at the somatic membrane using conventional and dynamic patch-clamp recording techniques, which enable the ciliary mechanism to be bypassed. A conditioning stimulus with an odorant-induced current markedly shifted the response range of action potentials induced by the same test stimulus to higher concentrations of the odorant, indicating odor adaptation. This effect was inhibited by charybdotoxin and iberiotoxin, Ca2+-activated K+ channel blockers, suggesting that somatic Ca2+-activated K+ currents regulate odor adaptation by modulating spike encoding. I conclude that not only the ciliary machinery but also the somatic membrane currents are crucial to odor adaptation.     

SENSORY INTERACTIONS IN MIXTURES

Psychological studies have assessed the intensity of simple sensory mixtures, both in taste and olfaction. In taste mixtures, suppression or partial masking among the components is often observed. An analogous result is often found in odor mixtures, counteraction of one component in the presence of a second odor. These effects, particularly taste suppression, are also observed in food systems. Interactions between sensory modalities are far more complex, ranging from inhibition of taste and odor sensations by trigeminal irritation, to relative independence of tastes from odor stimulation and independence of odors from tastes.     

THE ROLE OF THE KNOWLEDGE OF COLOR AND BRAND NAME ON THE CONSUMER'S HEDONIC RATINGS OF TOMATO PUREES

The objective of study was to assess the influence of the knowledge of color and brand name on consumer response to tomato purees. From focus group interviews it was found that taste and odor of fresh tomato were the most important drivers of consumer choice. To quantify that information a questionnaire was developed and it was submitted to fifty consumers of tomato puree. The questionnaires answers confirmed those coming from the focus group, although same discrepancies were found with those derived from the consumer test. Results showed that color and brand name seemed to be more important than taste and odor of fresh tomato, in fact those attributes, when consumers were tasting samples, affected not only hedonic judgments but sensory perception as well. Finally physico-chemical parameters derived from instrumental measurements have been compared to sensory data by means of PCA in order to validate the consumers'assessment of sensory attributes of tomato puree.     

Regulatory processes of hunger motivated behavior

While food intake and body weight are under homeostatic regulation, eating is a highly motivated and reinforced behavior that induces feelings of gratification and pleasure. The chemical senses (taste and odor) and their evaluation are essential to these functions. Brainstem and limbic glucose-monitoring (GM) neurons receiving neurochemical information from the periphery and from the local brain milieu are important controlling hunger motivation, and brain gut peptides have a modulatory role on this function. The hypothalamic and limbic forebrain areas are responsible for evaluation of reward quality and related emotions. They are innervated by the mesolimbic dopaminergic system (MLDS) and majority of GM neurons are also influenced by dopamine. Via dopamine release, the MLDS plays an essential role in rewarding-reinforcing processes of feeding and addiction. The GM network and the MLDS in the limbic system represent essential elements in the neural substrate of motivation.     

A conditioned aversion study of sucrose and SC45647 taste in TRPM5 knockout mice

Previously, published studies have reported mixed results regarding the role of the TRPM5 cation channel in signaling sweet taste by taste sensory cells. Some studies have reported a complete loss of sweet taste preference in TRPM5 knockout (KO) mice, whereas others have reported only a partial loss of sweet taste preference. This study reports the results of conditioned aversion studies designed to motivate wild-type (WT) and KO mice to respond to sweet substances. In conditioned taste aversion experiments, WT mice showed nearly complete LiCl-induced response suppression to sucrose and SC45647. In contrast, TRPM5 KO mice showed a much smaller conditioned aversion to either sweet substance, suggesting a compromised, but not absent, ability to detect sweet taste. A subsequent conditioned flavor aversion experiment was conducted to determine if TRPM5 KO mice were impaired in their ability to learn a conditioned aversion. In this experiment, KO and WT mice were conditioned to a mixture of SC45647 and amyl acetate (an odor cue). Although WT mice avoided both components of the stimulus mixture, they avoided SC45647 more than the odor cue. The KO mice also avoided both stimuli, but they avoided the odor component more than SC45647, suggesting that while the KO mice are capable of learning an aversion, to them the odor cue was more salient than the taste cue. Collectively, these findings suggest the TRPM5 KO mice have some residual ability to detect SC45647 and sucrose, and, like bitter, there may be a TRPM5-independent transduction pathway for detecting these substances.