Amiloride Effects on Taste Quality: Comparison of Single and Multiple Response Category Procedures

Although there is compelling evidence that amiloride reducesthe intensity of Na⁺ and Li⁺ salts in humans, its effects onsaltiness are conflicting. Many salts elicit not only a saltytaste but also one or more side tastes (sweetness, sournessor bitterness). Some studies have shown a suppression of saltinessby amiloride; others show no effect on saltiness but a significantreduction in sourness. In the experiments demonstrating a reductionof saltiness, subjects estimated only saltiness; in those showingan amiloride effect on sourness and not saltiness, subjectsestimated all qualities on each trial. The present study examinesthe role of the psychophysical method in these conflicting results.We have investigated the effects of amiloride on taste qualityby modifying only the instructions to the subjects, keepingall other variables constant. One group of subjects (intensity-only)gave magnitude estimates of the overall intensity of a LiCIconcentration series. A second group (salty-only) was instructedto estimate the saltiness of the stimuli, and a third group(sour-only) estimated their sourness. Finally, a fourth group(profile) rated all of the taste qualities on each stimuluspresentation, using a modified taste profile method. The ratingsof all groups were made comparable by the use of 0.1 mM quinine-HCIas a modulus. When subjects used only one response category,amiloride reduced their estimates (of intensity, saltiness orsourness), but if subjects attended to all four qualities, amiloridespecifically reduced the sourness of LJCI and had no significanteffect on its saltiness. Comparison of the saltiness estimatesof the salty-only group to the sum of the salty and sour estimatesof the profile group demonstrated that subjects combined thesesensations when presented with only one response alternative.To reveal the effect of amiloride on a specific quality of asalt, the psychophysical method must allow subjects to attendto all qualities on each trial. These data and previous resultssuggest that apical Na⁺ channels on the taste receptor cellmembrane mediate the sourness but not the saltiness of Na⁺ andLi⁺ salts. Chem. Senses 22: 267–275, 1997.     

REGIONAL VARIATION IN SWEET SUPPRESSION

Differences in the sweet‐blocking efficacy of 2‐(4‐methoxyphenoxy) propanoic acid (PMP) for different sweeteners (sucrose and aspartame) and for various exposure areas of the mouth were found. Twenty participants rated sweetener solutions with and without PMP for sweetness, sourness, saltiness, bitterness and umami for stimulation of anterior tongue, posterior tongue and whole‐mouth areas. For sweetness ratings, suppression was significant for all stimulation areas. In the presence of PMP, stimulation of the posterior tongue yielded significantly higher sweetness ratings than stimulation of the anterior tongue for aspartame but not for sucrose. Sourness and bitterness ratings were significantly higher for anterior tongue than posterior tongue stimulations for aspartame but not for sucrose. The increases in sourness ratings in the presence of PMP were likely because of the sour taste PMP has at the concentration used. Results imply a difference between the front and the back of the tongue in the mechanisms involved in the perception of sweetness.     

Confusing tastes and smells: how odours can influence the perception of sweet and sour tastes

This study investigated the relationship between perception of an odour when smelled and the taste of a solution to which the odour is added as a flavorant. In Experiment 1 (E1) sweetness, sourness, liking and intensity ratings were obtained for 20 odours. Taste ratings were then obtained for sucrose solutions to which the odours had been added as flavorants. Certain odours were found to enhance tasted sweetness while others suppressed it. The degree to which an odour smelled sweet was the best predictor of the taste ratings. These findings were extended in Experiment 2 (E2), which included a second tastant, citric acid, and employed four odours from E1. The most sweet smelling odour, caramel, was found to suppress the sourness of citric acid and, as in E1, to enhance the sweetness of sucrose. Again, odours with low sweetness suppressed the sweetness of tasted sucrose. The study demonstrated that the effects of odours on taste perception are not limited to sweetness enhancement and apply to sour as well as sweet tastes. The overall pattern of results is consistent with an explanation of the taste properties of odours in terms of prior flavour-taste associations.     

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.     

Sensory integration in citric acid/sucrose mixtures

The scale values of perceived sweetness, sourness and totaltaste intensity of unmixed sucrose, unmixed citric acid andseveral citric acid/sucrose mixtures were assessed, using afunctional measurement approach in combination with a two-stimulusprocedure. The data showed that the scale values obtained werelinear with perceived taste intensity. It was demonstrated thatcitric acid suppresses the sweetness of sucrose and that, inversely,sucrose suppresses the sourness of citric acid. However, thissuppressive effect was not symmetrical in the range of concentrationsused. While the degree of sweetness suppression depended onlyon the citric acid level, the degree of sourness suppressiondepended on the sucrose as well as on the citric acid concentration.With regard to the perceived total taste intensity of citricacid/sucrose mixtures, it was shown that the sum of sweetnessand sourness approximately equals the total taste intensity.The implications of the present findings for the analytic—syntheticcontroversy and for taste interaction theories are discussed.     

Intensity and hedonic judgments of taste mixtures: an information integration analysis

Anderson's (1981) information integration approach was usedto examine taste mixture integration for intensity and hedonicjudgments of sucrose/sodium chloride and sucrose/citnc acidsolutions. In Experiment I, total intensity and hedonic ratingswere made for factorial combinations of sucrose and sodium chlorideor citric acid. The total intensity judgments produced an integrationpattern characterized by extreme subadditivity at high soluteconcentrations. (Subadditivity refers to the tendency for totalmixture intensity to be rated as less than the sum of the unmixedcomponent intensities.) The intensity judgment integration patternswere essentially identical for the two mixture types. However,the pattern of integration for the sucrose/sodium chloride andsucrose/citric acid mixtures differed for the hedonic ratings.Sucrose tended to eliminate the unpleasantness associated withincreasing concentrations of citric acid, while it only modulatedthis trend for sodium chloride. In Experiment II, subjects ratedthe individual sweet, salty and sour components of the mixturesto determine whether mixture suppression of the component tastescould account for the subadditivity of the total intensity judgmentsand/or the pattern of results for the hedonic ratings. It wasfound that sucrose suppressed the sour component of the sucrose/citricacid mixtures more than the salty component of the sucrose/sodiumchloride solutions This difference in component suppressionseemed to account for the hedonic integration patterns of thetwo mixture types which suggests that mixture suppression isan important factor to consider when predicting the pleasantnessof simple taste mixtures.     

Interactions between oral chemical irritation, taste and temperature

The oral chemical irritant, capsaicin, at 2, 4 and 8 p.p.m.,was combined in mixtures with sucrose (Experiment 1), sodiumchloride (Experiment 2) and soup (Experiment 3), each evaluatedat two temperatures. These mixtures were rated for their sweetnessand/or saltiness, intensity of burning sensation and total mixtureintensity. In both solution and soup, sweetness was suppressed,whereas saltiness showed only minor suppression in low NaCl,high capsaicin mixtures. The burning sensation produced by capsaicinwas uninfluenced by sucrose, while NaCl increased the burningsensation. Total mixture intensity was entirely determined bycapsaicin concentration in mixtures with sucrose, although NaClcontributed in NaCl/capsaicin mixtures. Varying temperatureinfluenced the burning sensation and total intensity of sucrose/capsaicinmixtures, but did not modulate the effects of capsaicin on taste.Explanations of taste suppression in terms of cognitive andstructural models are examined. The differential effect of capsaicinon sweetness and saltiness is also considered in terms of theirritant properties of NaCl.     

SOURNESS–SWEETNESS INTERACTIONS IN DIFFERENT MEDIA: WHITE WINE,ETHANOL AND WATER*

The aim of this work was to study the sourness–sweetness interactions in water, white wine and alcoholic environment to interpret sweet/sour perception in low concentrations within the range normally encountered in white wine. Nine trained assessors rated sweetness and sourness intensity in mixtures of fructose (11.1, 25.0 and 38.9 mM) and tartaric acid (pH 3.0, 3.4 and 3.8) in water and wine (experiment 1) or ethanol solutions at 2.0, 4.0 and 12.0% v/v (experiment 2). The range of quantitative responses was larger for sourness than for sweetness in the three media. The global sourness intensity perception in wine mixtures was significantly lower than in water and ethanol mixtures, indicating the effect of other wine components. The suppressive effect of tartaric acid on fructose sweetness was stronger than the suppressive effect of fructose on tartaric acid sourness.     

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.     

An assessment of binary mixture interactions for nine sweeteners

Binary mixtures of acesulfame K, aspartame, sodium cyclamate,fructose, glucose, stevioside, sodium saccharin, sucrose andxylitol were assessed using factorial mixture designs. A simpleadditive model was used to generate predictions for the sweetnessof the mixtures and these predicted responses were comparedto the observed sweetness ratings of the mixtures. It was foundthat the mixtures tended to exhibit superadditivity at low concentrations,additivity at intermediate concentrations and subadditivityat high concentrations. Synergistic and suppressive effectsin the mixtures were evaluated by comparing mixture responsesto the sweetness ratings of ‘self-mixtures’. Self-mixturedata were generated by treating a mixture of a substance withitself as if it were a mixture of two different substances.Synergism was defined as a mixture response that was greaterthan the sweetness of the component self-mixtures, and suppressionwas defined as a mixture response that was less than the sweetnessof the component self-mixtures. Of the 31 binary mixtures studied,18 showed synergism, two showed suppression and 11 did not differsignificantly from their components. It is hypothesized thatmultiple sweetness receptors or release from bitter suppressionmay account for the synergistic effects.     

Enhancement of sucrose sweetness with soluble starch in humans.

The effect of soluble starch (acid-modified starch) on taste intensity was investigated in human subjects. Different concentrations of sucrose (Suc), six sweeteners, NaCl, quinine-HCl (QHCl) and citric acid (Cit) were dissolved in either distilled water (DW; standard) or starch solution (test solution). The solutions were presented to naive subjects and each subject was requested to taste and compare the sweetness intensity between the standard and test solutions based on a scale ranging from +3 (enhanced) to -3 (inhibited). A greater sweetness intensity occurred with Suc at different concentration (0.1-1.0 M) dissolved in soluble starch (0.125% to 4.0%) than with Suc in DW. Similarly, five other different products of soluble starch at 0.25 and 4.0% resulted in enhancement of sweetness for 0.3 and 1.0 M Suc. With the sole exception of the taste of 0.3 M Suc, sweet enhancement did not occur with 0.43 M fructose, 0.82 M glucose, 0.82 M sorbitol, 0.0037 M aspartame, 0.0042 M saccharin-Na or 0.016 M cyclamate. Neither the saltiness of NaCl (0.01-0.3 M), the bitterness of QHCl (0.00003-0.001 M) nor the sourness of Cit (0.0003-0.01 M) were affected by the soluble starch. These results suggest that the taste enhancing effects of soluble starch on Suc sweetness might depend not only on the taste transduction mechanism, but also on the molecular interaction between Suc and soluble starch.     

The addition of CO2 to traditional taste solutions alters taste quality

Previous studies of the effect of carbonation on taste perception have suggested that it may be negligible, manifesting primarily in increases in the perceived intensity of weak salt and sour stimuli. Assuming CO2 solutions in the mouth stimulate only trigeminal nerve endings, this result is not altogether surprising; however, there are neurophysiological data indicating that CO2 stimulates gustatory as well as trigeminal fibers. In that case, carbonation might alter the quality profile of a stimulus without producing substantial changes in overall taste intensity--much as occurs when qualitatively different taste stimuli are mixed. To address this possibility, subjects were asked to rate the total taste intensity of moderate concentrations of stimuli representing each of the basic tastes and their binary combinations, with an without added carbonation. They then subdivided total taste intensity into the proportions of sweetness, saltiness, sourness, bitterness and 'other taste qualities' they perceived. The addition of carbonation produced only small increases in ratings of total taste intensity. However, rather dramatic alterations in the quality profiles of stimuli were observed, particularly for sweet and salty tastes. The nature of the interaction is consistent with a direct effect of carbonation/CO2 on the gustatory system, although the possibility that at least some of the observed effects reflect trigeminal-gustatory interactions cannot be ruled out.      

A COMPARISON OF METHODS FOR MONITORING INDIVIDUAL PERFORMANCES IN TASTE SELECTION TESTS

The evaluation of panel performance was made by three methods: average of correct responses (A), comparison of distances of individual standardized judgments to the average standardized responses (D) and a principal components analysis (PCA). Thirty assessors identified water and basic tastes and discriminated different sweet stimuli in neutral or acidified vehicles using R‐index rating and ranking tests. By A and D methods 22 assessors were qualified as proficient. Composition of both panels was identical except for one judge. The output from PCA provided a graphical representation of the performance of the assessors and retained different subsets of 24–26 panelists for different proposals as discrimination of sweetness in acidified beverages, recognition of bitterness, sourness and discrimination of slight sweetness or evaluation of saltiness.     

Anterior Tongue Stiomulation with Amiloride Suppresses NaCl Saltiness, but not Citric Acid Sourness in Humans

Suppression of the saltiness of NaCI solutions by amiloride,a sodium channel blocker, has previously been reported a numberof times in humans. This suppression was seen with techniquesthat involved stimulation of small areas of the tongue. It wasnot certain, however, whether amiloride would suppress saltinesswith stimulation of a much larger area of the tongue; one publishedstudy, in fact, found negative results with whole mouth stimulation.For this study, eight subjects dipped a large part of the anteriorportion of the tongue into a 10-ml sample of NaCI solution,or a NaCI and amiloride solution, and reported its magnitudeof saltiness intensity. The results show that amiloride suppressedthe saltiness of NaCI when a large area of the anterior tonguewas stimulated. Consistent with previous studies, there wasindividual variability across subjects in this suppressive effectof amiloride. This study also used this method to test the effectsof amiloride on the sourness of citric acid, which was not expectedto be affected. No suppression of sourness was seen with amiloride.Chem. Senses 21: 113–120, 1996.     

Effect of sweetener type and of sweetener and acid levels on temporal perception of sweetness, sourness and fruitiness

Sweetness, sourness and fruitiness of 18 orange-flavored solutions,with three levels of citric acid (0.75, 1.5, 2.25 g/1) and threeequi-sweet levels of either sucrose (80, 100, 120 g/l) or aspartame(0.6, 0.7, 0.8 g/l), were evaluated by time–intensitymethodology. At these concentrations, a larger range in sournessintensity than in sweetness was produced, resulting in greatersuppression of sweetness by increasing acid levels than of sournessby increasing sweetener levels. Although aspartame samples hada longer duration of sweetness and fruitiness, sucrose and aspartamedid not interact differently with the sourness of citric acid.Fruitiness intensity and duration was enhanced by both sweetnessand sourness, but to a greater extent by sourness. Whether thisenhancement is attributable to a cognitive association of sweetnessor sourness with fruitiness or is due to the inability of thesubjects to separate sweet and sour tastes from orally perceivedfruity flavor cannot be concluded from this study.     

Space and time separation of taste mixture components

Under natural conditions, all component qualities of taste mixturesco-exist in space and time on the tongue surface. The significanceof space-time coherence of taste sensations was investigatedby separating mixture components on the tongue. First, it wasshown that a mixture of sucrose plus sodium chloride is judgedto have a qualitatively different sweet taste from a mixtureof sucrose plus quinine sulphate when all taste qualities aretogether in space and time. This occurred even though mixtureswere matched for sweetness intensity, and the saltiness of theformer matched the bitterness of the latter. When, however,mixture components were spatially and temporally separated,the judged quality differences were eliminated. When the componentswere separated in space only, quality difference judgments werereduced in magnitude, but not eliminated. The results are consistentwith the hypothesis that space-time coherence of taste sensationsresult in perceptual blending and a consequent failure of selectiveattention to any single component. It is suggested that tastemixtures can be considered as single tastes even though componentqualities can be reliably identified using other techniques.     

Contextual effects in the perception of quinine HCl/NaCl mixtures

Stimulus distribution, stimulus spacing and stimulus range affectmean intensity ratings of solutions of unmixed tastants. Thepresent study compares contextual effects for mixture ratingswith those for unmixed stimuli: if ratings for mixed and unmixedstimuli are differentially affected by context, the degree ofmixture suppression inferred from the responses is context-dependent.Bitterness intensity ratings for unmixed quinine and quinineHCl/NaCl mixtures were not differentially affected by shiftsin stimulus frequency distribution. The subjects' tendency tobe consistent in their responses to identical stimuli, resultsin a transfer of stimulus context of previous sessions to thenext.     

Effect of repeated presentation on sweetness intensity of binary and ternary mixtures of sweeteners

The purpose of the present study was to determine the effect of repeated presentation of the same sweet stimulus on sweetness intensity ratings. The sweet stimuli tested in this study were binary and ternary blends of 14 sweeteners that varied widely in chemical structure. A trained panel evaluated the sweetness intensity over four sips of a given mixture presented at 30 s intervals. The individual components in the binary sweetener combinations were intensity-anchored with 5% sucrose, while the individual sweeteners in the ternary mixtures were intensity-anchored with 3% sucrose (according to formulae developed previously). Each self-mixture was also evaluated (e.g. acesulfame-K-acesulfame-K). The main finding of this study was that mixtures consisting of two or three different sweeteners exhibited less reduction in sweetness intensity over four repeated sips than a single sweetener at an equivalent sweetness level. Furthermore, ternary combinations tended to be slightly more effective than binary combinations at lessening the effect of repeated exposure to a given sweet stimulus. These findings suggest that the decline in sweetness intensity experienced over repeated exposure to a sweet stimulus could be reduced by the blending of sweeteners.