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.     

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.     

The capacity of humans to identify components in complex odor-taste mixtures

Despite the fact that humans experience mixtures of odors and tastes each time they eat, little is known of their capacity to detect the individual components of foods. To investigate this capacity, 43 subjects were trained to identify three odors and three tastes and were required to indicate which of these could be identified in stimuli consisting of one to six components. Although the odor and taste components of most binary mixtures were identified, subjects encountered substantial difficulties with more complex mixtures with only two components being identified in the four- to six-component mixtures. In general, tastes were more easily identified than smells and were the only stimuli identified in the five- to six-component mixtures. Several mechanisms are proposed to account for the poor identification of components.     

Synthesis of tastes other than the 'primaries': implications for neural coding theories and the concept of 'suppression'

It has been demonstrated that mixtures of ‘primary’tastes may be perceived as singular rather than as multiple(Erickson and Covey, 1980). This finding may be accounted forin two different and testable ways. First, suppression: oneof the tastes in the mixture may be suppressed by the other(i.e., in a mixture of A and B, A suppresses the taste of B);the prediction which follows from this hypothesis is that asolution of A (the suppressing component) should be indistinguishablefrom the AB mixture. Secondly, synthesis: the singularity ofthe mixture may arise from the formation of a new taste, andin this case both components should be distinguishable fromtheir mixture. The present data indicate that the latter istrue in some cases. Since the taste of these mixtures must beother than the primary tastes used to form them, these findingsquestion the presumption that only four ‘primaries’are adequate to describe the range of tastes.     

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.     

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.     

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.     

Binary taste mixture interactions in prop non-tasters, medium-tasters and super-tasters

It is generally assumed that the mutual, but asymmetric, suppression of the components in binary taste mixtures is an invariant property of the human psychophysical response to such mixtures. However, taste intensities have been shown to vary as a function of individual differences in sensitivity, indexed by the perceived bitterness of 6-n-propylthiouracil (PROP). To determine if these variations in taste perception influence taste mixture interactions, groups of PROP super-, medium- and non-tasters assessed four binary taste mixtures: sweet-bitter [sucrose/quinine hydrochloride (QHCl)], sweet-sour (sucrose/citric acid), salty-bitter (NaCl/QHCl) and salty-sour (NaCl/citric acid). In each experiment, subjects received factorial combinations of four levels of each of two tastants and rated individual taste intensities and overall mixture intensity. For each taste quality, super-tasters typically gave higher ratings than either medium- or non-tasters, who tended not to differ. There were also group differences in the interactions of the mixtures' components. Super-tasters rated the overall intensity of the mixtures, most likely reflecting integration of the taste components, as greater than medium- and non-tasters, who again showed few differences. In sweet-bitter mixtures, non-tasters failed to show the suppression of sweetness intensity by the highest QHCl concentration that was evident in super- and medium-tasters. These data show that the perception of both tastes and binary taste mixture interactions varies as a function of PROP taster status, but that this may only be evident when three taster groups are clearly distinguished from one another.     

Perceptual interactions between characteristic notes smelled above aqueous solutions of odorant mixtures

Twenty-two experienced panelists rated odor intensity of aqueous solutions of citral, octen-1-ol-3, and hexanal. The panel assessed unmixed components and mixtures (9 binary and 4 ternary). In sensory sessions dedicated to mixtures (n = 6), evaluation was focused on one target odor, presented at a fixed concentration. All components had lower odor intensity on mixed presentations. In many cases, information obtained from simpler systems was not extended to complex mixtures. In a mixture, the competition between odorant molecules on qualitative aspects (dominance/suppression) imbalanced components contribution, anticipated from the quantitative distribution. Hexanal appeared to be the potentially weaker odorant in paired combinations, whereas octen-1-ol-3 had a lower relative impact on ternary systems. Suppression of the odor of octen-1-ol-3 and a concomitant increase in the odor of hexanal was common to all ternary mixtures. Reciprocal inhibition of octen-1-ol-3 and citral odors through perceptual interactions was suspected. Mutual suppression is suspected to have eased the perception of hexanal intensity.     

PERCEPTION OF TASTE MIXTURES BY EXPERIENCED AND NOVICE ASSESSORS1

The total intensity, sweetness, and acidity of sucrose/citric acid mixtures were judged by two types of taste panel: experienced assessors, most of whom had had many years of experience in sensory evaluation; and novice assessors, none of whom had previously taken part in a taste experiment. In other respects the experimental conditions remained almost constant. There was good correspondence between the two panels, particularly for judgments of total intensity, indicating that novice and experienced assessors evaluate taste mixtures in the same way. However, there was also an indication that experience on sensory panels may attenuate taste suppression, the suppression of acidity by sweetness being less pronounced for the experienced panel than for the novice panel. The implications for mixture perception are noted.     

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.     

The taste, odor and hedonic quality of polyglycerols

Twenty subjects judged the taste and odor intensity and thetaste and odor pleasantness/unpleasantness of five concentrationsof sucrose, glycerol, a commercial triglycerol, a syntheticlinear diglycerol and a synthetic linear triglycerol. Judgmentsof intensity were made using the method of magnitude estimation;judgments of pleasantness/unpleasantness were made using a graphicline scale. Only the two linear polyglycerols had appreciableodor intensity. Both were described as having an ‘acrid’or ‘burnt caramel’ quality. The odor exponent forthe linear triglycerol was extremely high (1.44) and may beattributed to its intensely unpleasant quality. Sucrose wascharacterized solely by sweet taste, glycerol and the commercialtriglycerol by sweet and bitter tastes, the linear diglycerolby sweet, bitter and sour tastes, and the linear triglycerolby bitter and sour tastes. The relationships between perceivedtaste intensity and concentration were well described by powerfunctions, although the slope of the psychophysical functionfor the linear triglycerol was markedly lower than that forthe other compounds. The relative order of taste intensitieswas: linear triglycerol > sucrose > glycerol = lineardiglycerol > commercial triglycerol. Judgments of taste (andodor) pleasantness/unpleasantness showed only sucrose and glycerolto have positive hedonic qualities. All the polyglycerols werejudged unpleasant at all concentrations. Differences in thetaste and odor characteristics of the commercial and synthetictriglycerols were attributed to the commercial product beinga mixture of over 20 compounds. Although the synthetic lineardi- and triglycerols are effective in lowering water activity,these data suggest that more purified crystalline forms mustbe synthesized before they can be used effectively as humectantsfor intermediate moisture foods.     

Decline in taste and odor discrimination abilities with age, and relationship between gustation and olfaction

It is important to learn about changes in both taste and odor perceptions with increasing age, because the taste of foods we encounter in our daily life is strongly affected by their smell. This study discusses the difference in qualitative taste and odor discrimination between the elderly and the young. Tastants and odorants used in this study were presented not as single stimuli but as a taste mixture (sucrose and tartaric acid) and an odor mixture (beta-phenylethyl alcohol and gamma-undecalactone). The results showed that quality discrimination abilities of the elderly subjects for both taste and odor were significantly lower than those of the young subjects, indicating a decline in quality discrimination abilities related to age. Also, a moderate but significant correlation was observed between the taste discrimination ability and the odor discrimination ability. We measured thresholds for single-taste and odor components in mixtures and compared them between the elderly and the young to investigate the cause for these findings.     

Reaction time in the perceptual processing of taste quality

A series of experiments examined possible strategies human subjectsmight use in tasks requiring rapid detection or recognitionof a taste quality. In a reaction time (RT) paradigm, subjectswere to decide whether each of a series of stimuli flowed overthe tongue contained a previously designated target taste. Severaltasks of varying difficulty were used. The simplest task requiredsubjects to decide whether the target taste or water was presented.The most difficult task required discrimination between twodifferent target tastes in a series of mixtures formed by orthogonallycombining the target taste with two different irrelevant tastes.The speed at which subjects could detect and/or recognize targettastes was related to the RT for the particular taste. However,it was also clear that other variables, including the specificstimuli in the mixtures and the cognitive demands placed onthe subjects, influenced performance. These results suggestthat differences in taste onset time, as indexed by RT, canserve as a cue which subjects use to aid identification of singletastes in a mixture. It is concluded that the ease with whichsubjects can identify single tastes in a mixture is relatedto, among other variables, the differences in taste onset timebetween the tastes.     

Taste Responses to Binary Mixtures of Amino Acids in the sea catfish, Arius felis

In vivo electrophysiological recordings in the sea catfish,Arius felis, showed that the magnitude of the integrated facialtaste responses to binary mixtures of amino acids was predictablewith knowledge obtained from previous cross-adaptation studiesof the relative independence of the respective binding sitesof the component stimuli. Each component from which equal aliquotswere drawn to form the mixtures was adjusted in concentrationto provide for approximately equal reponse magnitudes. The magnitudeof the taste responses to binary mixtures whose component aminoacids showed minimal cross-adaptation was significantly greaterthan that to binary mixtures whose components exhibited considerablecross-reactivity. There was no evidence for mixture suppression.The relative magnitude of the taste responses in the sea catfishto stimulus mixtures is similar to that previously reportedfor olfactory receptor responses in the freshwater channel catfishand chorda tympani taste responses in the hamster. Chem. Senses21: 45–53, 1996.     

Modality-specific neural effects of selective attention to taste and odor

The insular cortex is implicated in general attention and in taste perception. The effect of selective attention to taste on insular responses may therefore reflect a general effect of attention or it may be (taste) modality specific. To distinguish between these 2 possibilities, we used functional magnetic resonance imaging to evaluate brain response to tastes and odors while subjects passively sampled the stimuli or performed a detection task. We found that trying to detect a taste (attention to taste) resulted in activation of the primary taste cortex (anterior and mid-dorsal insula) but not in the primary olfactory cortex (piriform). In contrast, trying to detect an odor (attention to odor) increased activity in primary olfactory but not primary gustatory cortex. However, we did identify a region of far anterior insular cortex that responded to both taste and odor "searches." These results demonstrate modality-specific activation of primary taste cortex by attention to taste and primary olfactory cortex by attention to odor and rule out the possibility that either response reflects a general effect of attentional deployment. The findings also support the existence of a multimodal region in far anterior insular cortex that is sensitive to directed attention to taste and smell.     

SENSORY ADAPTATION

Sensory evaluation makes use of the remarkable virtuosity and range of the human senses as a multi-purpose instrument for measuring the sensory characteristics of foods. The brain protects itself from an overload of information from the senses by two processes: feature extraction and adaptation. The former involves information reduction by the extraction of selected features from the environment; these form the basis for the reconstruction of the percept in consciousness. The latter, adaptation, involves the attenuation of repetitive and constant input so as not to overload the brain with redundant information.
The effects of adaptation can be observed for all senses. For the chemical senses, the effect is that a constant odor or taste stimulus will be perceived as decreasing in intensity while sensitivity to that stimulus is also decreased. For sensory evaluation, this poses problems. It means that a taste or odor has a tendency to vanish while it is being observed and that sensitivity to subsequent stimuli will be altered. Such sensitivity drift in the human instrument must be anticipated in the design of measurement procedures for the sensory evaluation of food.
For taste, adaptation changes caused by the measurement method can be seen to be largely responsible for disagreements in the literature concerning threshold and intensity measurement. Adaptation is also a contributing factor, but by no means the only one, in determining the relative discriminability of sensory difference tests. It is worth noting, however, that adaptation is not always a disadvantage; it can sometimes be used to advantage in sensory testing procedures.     

Application of the U and gamma' models in binary sweet taste mixtures

The U and Gamma' models of sensory interactions, successfully applied in olfaction for several years, are tested here using data from published studies on sweetness. The models are subsequently tested on new data obtained in studies of binary mixtures of four sodium sulfamates. The U model allows for the estimation of a global interaction, whereas the Gamma' model allows for the distinction between that which is due to an intrinsic interaction in the mixture itself and that which may be due to the power function exponents in the mixture. The models give satisfactory predictions for observed phenomena of sweet taste suppression, synergism or pure additivity. Additionally, they appear to be more suitable than other models recently applied in taste, particularly the equiratio model. Application of the models to the sulfamate mixtures, reveals additivity for sodium cyclohexylsulfamate (cyclamate)/potassium cyclohexylsulfamate and sodium cyclohexylsulfamate/sodium exo-2-norbornylsulfamate, respectively; whereas for sodium cyclohexylsulfamate/sodium 3-bromophenylsulfamate, the models revealed a slight hypo addition which is simply due to the dissimilarity values of the power function exponents of the components.     

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.     

THE TASTES OF ARTIFICIAL SWEETENERS AND THEIR MIXTURES

0s scaled the taste intensity (bitterness, sweetness) of artificialsweeteners, mixtures of artificial sweeteners, and glucose.Sweetness of glucose conformed to a power function, whereasneither sweetness of artificial sweeteners nor their bitternessdid. The total taste intensity of mixtures was often lower thanthe taste intensities of the components, suggesting suppression,although in many instances the suppressive effects disappearedat high concentrations.