Taste Mixtures Research Articles

Identification of the components of taste mixtures by adults and children.

Recently, it was reported that cognitive factors may have impeded the analysis of binary taste mixtures by 8- to 9-year-old children, resulting in them identifying only the strongest component in each mixture compared to adults who identified both components (Oram, Laing, Freeman, & Hutchinson, 2001). The current study aimed to overcome this limitation by employing different training procedures and different instructions when conducting the task. In particular, the training involved familiarization with the two stimuli at several levels of perceived intensity to facilitate identification of the varying levels of each stimulus in mixtures. Using a selective attention procedure, adults and children were required to identify the components of stimuli that contained water, sucrose (sweet), sodium chloride (salty), or were binary mixtures of these two tastants. Both groups were successful in identifying the component(s) in each stimulus, demonstrating the importance of appropriate training and test procedures when studying the sensory responses of children. Together with data from the earlier study by Oram et al. (2001) it is concluded that 8- to 9-year-olds have the cognitive skills to analyze taste mixtures in terms of the identity of the components and their perceived strengths.

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.

Counter-conditioning Following Human Odor–Taste and Color–Taste Learning

Tasting an odor in citric acid can result in that odor becoming less liked and sourer smelling. Both these changes appear to be resistant to extinction. As acquired hedonic responses to faces are also resistant to extinction, but amenable to counter-conditioning, the current experiment tested whether conditioned changes in odors could also be counter-conditioned. Presenting an odor in citric acid and then in sucrose did not change the perceptual and hedonic consequences of the initial pairing in citric acid. However, pairing a color with citric acid and then with sucrose produced a color–sweetness association. These findings are suggestive of a difference in counter-conditioning sensitivity between odors and other types of stimuli, which may result from encoding odor–taste mixtures in a relatively unanalyzed configural form.

The across-fiber pattern theory and fuzzy logic: A matter of taste

This article discusses a Fuzzy Logic (FL)-based model of neural coding and integration, proposed to be a formal extension of the Across-Fiber Pattern (AFP) theory. FL integration is conceptually similar to Bayesian reasoning, thus providing close-to-optimal decisions, and is also robust in that it does not require complete information. As a formal extension of AFP theory, the FL model describes sensory integration given multiple sources of information. When applied to gustation, the FL model is suggested to describe integration of information at the level of real-time pattern of single neural responses, population coding, and taste perception, as well as to provide a suitable description of taste mixtures.

Taste in the Monkey Cortex

The sense of taste in humans differs substantially from that of rodents, from which a preponderance of gustatory electrophysiology derives. To establish a more appropriate neural model for human gustation, we recorded the activity of single neurons in the primary taste cortex in 11 alert cynomolgus macaques. Taste cells composed 6% of all neurons encountered. Another 24% responded during mouth and jaw movements, and 4% were sensitive to tactile stimulation of the mouth. Smaller numbers responded during olfactory or visual stimulation, or when the monkey extended his tongue. Taste cells could be divided into four statistically independent groups, corresponding to those most responsive to glucose (38%), NaCl (34%), quinine (22%), or HCl (5%). The location of a taste cell did not predict its response profile, i.e., there was no clear topographic organization of taste sensitivity. We established neural thresholds and intensity-response functions to the basic stimuli and determined that—with the exception of HCl, to which the macaque is relatively insensitive—they were similar to those reported by human subjects. We then turned to the coding of taste quality, as inferred in macaques from the patterns of neural activity elicited by each of greater than 100 stimuli. The results proved generally faithful to human reports of the perceived qualities of these same tastants. Finally, an investigation of taste mixtures revealed a degree of mixture suppression and interaction among basic qualities similar to those reported by humans. We conclude that the alert macaque offers a reliable neural model for human gustation.

Perceived intensity and persistence of heterogeneous taste mixtures

Perceived intensity and persistence of single taste stimuli, binary and ternary heterogeneous taste mixtures in a sweet acidulated system were assessed. The effects of a sweetener, aspartame (APM) on sourness, and of citric acid (CA) buffered with different levels of its sodium salt (NaCit) on sweetness were examined. A sensory panel scored intensity and duration of single and mixed stimuli formulated with APM (0, 1, 2 or 4 mM), CA (50 mM) and NaCit (0, 10, 20 or 40 mM). The analysis of frequency of qualities assigned by subjects depicts a great similarity in the intensity as well as in the persistence attributes. All stimuli were perceptually measurable into the four taste qualities, but sour and sweet were the most frequent responses obtained when tasting solu tions. Perceived sourness dropped by adding NaCit or APM to the acid solution. Sour persistence showed roughly equivalent results. The intensity and duration of sweet quality rose with APM concentration. A drop in the pH of the solutions diminished sweet persistence but not intensity scores. These data suggested the operation of similar mechanisms for processing intensity and persistence in the present conditions.

Detection of Very Complex Taste Mixtures: Generous Integration Across Constituent Compounds

Mixtures of compounds can often be tasted even when all of their components are too weak to be tasted separately. Such mixtures are said to be integrative. Integration was demonstrated by mixing compounds in concentrations proportional to their separate detection thresholds and then measuring the detection threshold of the mixture as a whole by forced choice with plain water. Mixtures of 3, 6, 12, and 24 compounds were thus evaluated. With earlier data on two-, three-, and four-component mixtures (Stevens, J. C. Detection of taste in mixture with other tastes: Issues of masking and aging. Chem. Senses 21: 211–221; 1996.), the results show that the concentration of any constituent compound goes down in approximate proportion to the number of compounds with which it is in mixture. This nearly complete integration seems to describe mixtures of like-quality compounds, of unlike-quality compounds, and of both like- and unlike-quality compounds. Integrative mixtures of the sort studied here provide a model for the detection of the ultracomplex stimuli of everyday life, such as foods and drinking waters. Although the degree of integration may trail off slightly with mixtures of high complexity, the present result proffers no limit on the number of compounds that can be at least partially integrated. In principle, integration permits the detection of natural substances whose myriad components could all be far below threshold. The mechanism of taste integration is speculative, but the facts are congenial to the hypothesis of multiple parallel channels for the processing of intensity and quality.

ELIMINATING ARTIFACTS IN THE STUDY OF SINGULARITY/MIXEDNESS OF TASTE STIMULI

ABSTRACTTaste stimuli with a single solute were not judged as tasting equally singular; this confirmed earlier results, except in this study the possibility of the results being caused by trigeminal effects was virtually eliminated. Binary mixtures were generally perceived as more mixed than single solute stimuli, indicating a degree of analytic mixing. However, a caffeine/KCl mixture was perceived as more singular tasting than some single solute stimuli, indicating a degree of synthetic mixing. A preliminary examination was made of the hypotheses that the perceived mixedness of a taste mixture might be determined by a summation of the mixedness of its components or alternatively by latency effects.

Responses of the hamster chorda tympani nerve to binary component taste stimuli: evidence for peripheral gustatory mixture interactions

Studies of taste mixtures suggest that stimuli which elicit different perceptual taste qualities physiologically interact in the gustatory system and thus. are not independently processed. The present study addressed the role of the peripheral gustatory system in these physiological interactions by measuring the effects of three heterogeneous taste mixtures on responses of the chorda tympani (CT) nerve in the hamster ( Mesocricetus auratus). Binary taste stimuli were presented to the anterior tongue and multi-fiber neural responses were recorded from the whole CT. Stimuli consisted of a concentration series of quinine. HCI (QHCI: 1–30 mM), sodium chloride (NaCl: 10–250 mM). sucrose (50–500 mM) and binary combinations of the three different chemicals. Each mixture produced a unique pattern of results on CT response magnitudes measured 10 s into the response. Sucrose responses were inhibited by quinine in QHCI-sucrose mixtures. Neural activity did not increase when quinine was added to 50–250 mM NaCl in QHCI NaCl mixtures. However, the neural activity elicited by sucrose-NaCl mixtures was greater than the activity elicited by either component stimulus presented alone. The results demonstrate that gustatory mixture interactions are initiated at the level of the taste bud or peripheral nerve. Mechanisms for these interactions are unknown. The results are consistent with one component stimulus modifying the interaction of the other component stimulus with its respective transduction mechanism. Alternatively, peripheral inhibitory mechanisms may come into play when appetitive and aversive stimuli are simultaneously presented to the taste receptors.

Taste-mixture suppression: functional dissection of cellular and paracellular origins.

1. Chorda tympani (CT) nerve responses were recorded during simultaneous current and voltage clamping of the lingual receptive-field epithelium to examine the role of field potential in taste mixture suppression between sodium gluconate (NaG) and potassium gluconate (KG). 2. Under zero current-clamp conditions, CT responses to 100 mM NaG were suppressed by 63% when presented in mixture with 250 mM KG. At this concentration, KG alone elicited no measurable neural activity, but produced a large submucosal-positive field potential. 3. When CT responses to 100 mM NaG were obtained with voltage clamp at the zero-current clamp field potential of the NaG/KG mixture, they were suppressed by only 30% relative to NaG responses under zero-current clamp. Similarly, CT responses to the mixture of NaG and KG measured while voltage was clamped at the field potential of NaG alone were slightly elevated, but not to the magnitude of zero-current clamp responses to NaG. Therefore field potential-mediated suppression of CT responses to NaG accounts for only a part of the total mixture suppression between NaG and KG. 4. Analysis of the voltage dependence of CT responses to NaG indicated that the moderate field potential increase (8.9 mV) caused by the presence of KG in the mixture equates to a 43% increase in the apparent Km for NaG, from 110 to 157 mM. Use of this effective Km obviated the effect of field potential on CT responses to the NaG/KG mixture and permitted kinetic analysis of K+ blockade of Na+ responses. These analyses suggested that K ions block Na+ movement through apical Na+ channels in a voltage-independent manner with an apparent Ko of 405 mM. Importantly, direct inhibition of Na+ transduction by K+ can account for the part of mixture suppression not mediated by field potential. 5. These experiments reveal that mixture suppression between NaG and KG is derived from two distinct sources. Field potential, triggered largely by the limited mobility of both K+ and Na+ through taste bud tight junctions, globally modulates Na+ transduction. In addition, at the level of the apical Na+ channel, K ions directly block movement of depolarizing Na+ across taste receptor apical membranes.

Responses of primate cortical neurons to unitary and binary taste stimuli.

1. The responses of 126 neurons in primary gustatory cortices of two rhesus monkeys were recorded during sapid stimulation of the tongue with 18 taste stimuli. Ten of these stimuli were dissolved in distilled water (DW): 1.0 M sucrose (Suc), 0.1 M and 0.03 M sodium chloride (NaCl), 0.003 M hydrochloric acid (HCl), 0.001 M quinine hydrochloride (QHCl), 0.03 M monosodium glutamate (MSG), 0.03 M polycose, 0.3 M glycine, 0.1 M proline, and 0.1 M malic acid. Seven other stimuli were dissolved in 0.03 M MSG; the last stimulus was a mixture of 1.0 M Suc and 0.03 M NaCl. 2. The average spontaneous rate (2.2 +/- 0.2 spikes/s, mean +/- SE) and response to DW (2.5 +/- 0.2) of these 126 neurons was low but within the range previously reported for neurons in primate taste cortex. Suc was the most effective stimulus for 24.1% of the neurons tested followed by NaCl (15.7%), QHCl (14.8%), HCl (11.1%), MSG (10.2%), and other miscellaneous unitary gustatory stimuli (8.3%). Binary taste mixtures were the most effective stimuli for 15.7% of the sample. The net responses (corrected for DW, in spikes/s) for Suc-best (3.3), NaCl-best (4.3), HCl-best (3.4), QHCl-best (2.3), and MSG-best (4.1) were sluggish, but comparable with that reported previously. 3. The response breadth of the 82 neurons that responded best to either Suc, NaCl, HCl, or QHCl measured with the entropy coefficient indicated a moderate response breadth for these neurons (mean = 0.79; range = 0.30-0.98). According to the response criteria adopted in this experiment (water response +/- 1.96 SD), however, 81 of these 82 neurons (98.1%) responded to only one or two of the four basic taste stimuli. The disparity between the entropy- and criterion-based measures of response derive from the nature of the two statistics. Adjustments that would make the entropy statistic less inclusive and the definition of a response according to statistical criteria less exclusive would increase their concordance. 4. Three multivariate statistics (cluster, principal axis factor, and multidimensional analysis) were used to analyze the data. Cluster analysis enabled us to divide the 82 taste neurons into groups on the basis of response similarity. Each of the four largest groups was dominated by neurons that responded best to one of the four basic taste stimuli: Suc, NaCl, QHCl, and HCl (ranked in descending order); the fifth largest cluster contained neurons that responded best to MSG. Principal axis factor analysis demonstrated that 80.8% of the total variance could be accounted for by three factors. Neurons responding best to Suc, NaCl, and QHCl each were closely associated with one of those three factors, but the loadings of the HCl-best neurons were evenly distributed across all three factors. The communality coefficient of these three factors was > 80% for the Suc-, NaCl-, HCl-, and QHCl-best neurons; the MSG-best neurons, by comparison, had very few high loadings on any of these three factors and a correspondingly low communality coefficient of 40.4%, a difference that was statistically significant from the other four groups. Thus the three factors related to Suc-, NaCl-, HCl-, and QHCl-best neurons are not relevant to MSG-best neurons. We used multidimensional analysis to arrange the neurons that responded best to Suc, NaCl, HCl, QHCl, and MSG into five loosely arranged and partially overlapping clusters. A multidimensional space based on stimulus similarity showed that MSG was as different from the four basic taste stimuli as they were from one another. 5. Mixture suppression, a common observation in human psychophysical experiments, was examined at the neurophysiological level by including binary tastants in the stimulus battery. The average response of 19 Suc-best neurons to 1.0 M Suc (4.1 spikes/s) decreased to near 0 when the solvent was changed from DW to either 0.03 M MSG or 0.03 M NaCl. Similar decrements were observed in NaCl- and MSG-best neurons tested with Suc/NaCl mixtures.

Responses of single hamster parabrachial neurons to binary taste mixtures of NaCl with sucrose or QHCl.

1. Although human psychophysical responses to taste mixtures have been investigated extensively, there have been few reports on the neurophysiological coding of taste mixtures in the mammalian gustatory system. In recent studies we have investigated the responses of single third-order neurons in the hamster parabrachial nucleus (PbN) to anterior tongue stimulation with binary mixtures of heterogenous taste stimuli including sucrose+QHCl, sucrose+citric acid, and NaCl+citric acid. Some of these stimulus combinations evoked mixture suppression, or response frequencies that were less than that evoked by the more effective component (MEC) presented alone, which is analogous to the mixture suppression reported in human psychophysical studies of similar taste mixtures. In the current report we extend our investigation to include NaCl+QHCl and NaCl+sucrose mixtures. 2. The action potentials of single PbN neurons were recorded extracellularly. Four concentrations of each stimulus were employed: NaCl and sucrose at 0.001, 0.01, 0.1, and 1.0 M; QHCl at 0.00032, 0.0032, 0.032, and 0.1 M. All stimuli were tested alone and in mixture; the NaCl+sucrose and NaCl+QHCl mixtures were formed by pairing the four concentrations of each stimulus with the strongest concentration of the other stimulus. 3. For both NaCl+sucrose and NaCl+QHCl mixtures, the response frequencies evoked by the mixtures did not differ from those evoked by the MEC presented alone, whether averaged across all neurons or across subgroups of NaCl- or sucrose-best cells. Furthermore, the across-neuron patterns (ANPs) of mixture responses were similar to those of the MECs.(ABSTRACT TRUNCATED AT 250 WORDS)

Responses of single hamster parabrachial neurons to binary taste mixtures of citric acid with sucrose or NaCl.

1. Although taste experience generally arises from a mixture of gustatory stimuli, most neurophysiological studies of the mammalian central gustatory system have focused on responses to single chemical stimuli. Recently, in a study of single third-order neurons in the hamster parabrachial nucleus (PbN), we reported that mixture suppression occurs in the responses to binary mixtures of sucrose and QHCl presented to the anterior tongue. Mixture suppression was reflected both in reduced response frequencies and in an altered pattern of responses across neurons. In the current report we extend our investigation of CNS neuron responses to binary mixtures of heterogeneous stimuli to include sucrose+citric acid mixtures and NaCl+citric acid mixtures. The response to each mixture was compared with the response to the more effective component (MEC) presented alone, and those that differed by more than a selected criterion (based on response variability) were identified. 2. For all mixture responses recorded, 29% (79/256) involved mixture suppression (mixture response < MEC response), only 6% (18/276) were greater than the response to MEC, and 65% (179/276) did not differ from the response to the MEC. 3. In Experiments 1 and 2, neurons were tested with four concentrations of sucrose or citric acid each presented alone and in binary mixtures with a single strong concentration of the other stimulus. Sucrose suppression (mixture response < sucrose response) occurred in 24% of mixture responses and was exhibited almost exclusively by sucrose-best neurons, primarily to the mixtures that contained the stronger sucrose and citric acid concentrations. Sucrose suppression involved a 40% reduction of mixture response frequencies compared with responses to the sucrose component alone. 4. In Experiments 3 and 4, neurons were tested with four concentrations of NaCl or citric acid each presented alone and in binary mixtures with a single strong concentration of the other stimulus. NaCl suppression (mixture response < NaCl response) occurred in 21% of mixture responses and was displayed by both sucrose-best and NaCl-best neurons. NaCl suppression involved a 28% reduction in mixture response frequencies compared with responses to the NaCl component alone. In all experiments citric acid suppression (mixture response < citric acid response) was observed in only 6% of mixture responses and was relatively small in magnitude. 5. The across-neuron patterns (ANPs) of taste responses, which are correlated with behavioral measures of taste similarity, were compared for mixtures and components.(ABSTRACT TRUNCATED AT 400 WORDS)

Responses of single hamster parabrachial neurons to binary taste mixtures: mutual suppression between sucrose and QHCl.

1. Although taste experience typically arises from a mixture of gustatory stimuli, nearly all previous neurophysiological studies of the mammalian central gustatory system have focused on responses to single chemical stimuli. To begin to systematically examine CNS responses to taste mixtures, we recorded the extracellular activity of single third-order neurons in the hamster PbN to anterior tongue stimulation with binary mixtures of sucrose and QHCl. In experiment 1, neurons were tested with four concentrations of sucrose (0.001, 0.01, and 1.0 M) presented alone and mixed with 0.1 M QHCl. In experiment 2, neurons were tested with four concentrations of QHCl (0.00032, 0.0032, 0.032, and 0.1 M) presented alone and mixed with 1.0 M sucrose. 2. The response to each binary mixture was compared with the response to the more effective component (MEC) presented alone, and those that differed by more than a selected criterion (based on response variability) were identified. Of all mixture responses, 37% (59/158) involved mixture suppression (mixture response < MEC response), only 4% (6/158) were greater than the MEC, and 59% (94/158) were classified as not different than the response to the MEC. Most neurons that displayed mixture suppression did so at several mixture concentrations. 3. Sucrose suppression (mixture response < sucrose response) was prevalent among neurons most responsive to sucrose and for the mixtures that contained the stronger sucrose concentrations. Among neurons that displayed sucrose suppression, the magnitude of suppression was significantly correlated with sucrose response magnitude but not with QHCl response magnitude. These and other factors suggest that a neuron's capacity to display sucrose suppression to sucrose+QHCl mixtures is related to its sucrose sensitivity. 4. QHCl suppression (mixture response < QHCl response) was less prevalent than sucrose suppression, and the neurons that displayed QHCl suppression were almost exclusively a subset of those that displayed sucrose suppression to the same or different mixtures. This finding and the observation that one-third of all mixture responses involved mutual suppression (response to the mixture less than that to either component alone), suggest an association between the factors underlying sucrose suppression and QHCl suppression. 5. The across-neuron patterns (ANPs) of taste responses, which are thought to represent taste quality, were compared for mixtures and components. In general, the ANP for each mixture was similar to (significantly correlated with) the ANP of the more stimulatory component. However, for the mixture that evoked the greatest sucrose suppression, the mixture ANP was more similar to the ANP of the less stimulatory component.(ABSTRACT TRUNCATED AT 400 WORDS)

Evaluation of interaction in olfactory and taste mixtures

Models for the evaluation of interaction in olfactory and taste mixtures are compared to the isobole approach, which is widely used in other fields of bioraedical research. Analogies and differences are described in detail. The isobole approach has a profound theoretical basis and can be applied to all possible types of concentration-response relationships, in particular to all values of exponents in the widely used power function, and even if analytical expression of concentration—response relationships are not known. Due to this generality it leads to a substantial simplification of the evaluation procedures as compared to other methods used in taste and olfaction research. It can be applied to any number of components of a mixture. Response surface modeling and computer graphics is recommended in appropriate cases because it can provide information on the concentration dependence of interaction. Even though it has to be pointed out that there is no general consensus on the most appropriate approach for the evaluation of interaction between biologically active agents so far, we are led to the conclusion that good arguments can be made in supporting the application of the isobole method for taste and olfactory mixtures.

Contextual effects in taste mixture perception

Contextual effects in taste mixture perception

Mapping study of the parabrachial taste‐responsive area for the anterior tongue in the golden hamster

The locations of taste-responsive areas within the brainstem parabrachial nucleus (PBN), an obligatory taste relay in the golden hamster (Mesocricetus auratus), were mapped in relation to cytoarchitectural boundaries. The PBN was systematically searched for multiunit neural activity in response to a taste mixture composed of 0.1 M sucrose, 0.03 M NaCl, and 0.1 M KCl applied to the anterior tongue. Taste responses were located exclusively in one of three subdivisions of the medial PBN, which is thought to be specialized for gustatory processing, and in one of six subdivisions of the lateral PBN, which is thought to be specialized for general visceral processing. Based on Nissl-stained material, both the medial and lateral PBN subdivisions in the hamster were similar to those reported for the rat PBN. The largest group of taste-responsive cells encompassed two-thirds of the central medial subdivision, while a smaller group of taste cells was exclusively located within the ventral lateral subdivision. The two taste-responsive subdivisions are separated by the superior cerebellar peduncle and contain diverse cell types. The finding that anterior tongue taste may be exclusively represented in circumscribed cytoarchitecturally defined parts of two PBN divisions suggests that taste information from the anterior tongue is required for both specific gustatory and general visceral functions.

Perceptual integration of tertiary taste mixtures

Integration psychophysics was used to explore the taste perception of mixtures of sucrose, fructose, and citric acid. Three levels of each stimulus were varied in a 3 x 3 x 3 factorial design. Subjects rated total intensity, sweetness, and acidity of the 27 mixtures on graphic rating scales. Consistent with earlier work, the perceived total intensity of the tertiary mixtures was found to be dictated by the intensity of the (subjectively) stronger component alone (i.e., either the integrated sweetness or the acidity, whichever was the more intense). In contrast, the sweetness and acidity of the mixture were susceptible to mutual suppression: Sweetness suppressed acidity, acidity suppressed sweetness. There was, however, a difference between sucrose and fructose in their interactions with citric acid, fructose being the more susceptible to suppression. This selectivity of suppression indicates that the two sweetnesses could not have been inextricably integrated. Implications for taste coding are discussed, and the findings are reconciled in terms of two separate coding mechanisms: one for taste intensity, another for taste quality.

PERCEPTION OF TASTE MIXTURES BY EXPERIENCED AND NOVICE ASSESSORS1

ABSTRACT.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.

Temporal dissociation of taste mixture components

Temporal dissociation of taste mixture components