Sweetness intensity enhancement by pulsatile stimulation: effects of magnitude and quality of taste contrast

Upon stimulation with continuously alternating (pulsatile) taste concentrations, humans report higher average taste intensities than for continuous stimulation with the same average tastant concentration. We investigated the effect of the magnitude of concentration changes (concentration contrast) and the effect of taste quality changes (quality contrast) between alternating tastants on sweet taste enhancement. The perceived sweetness intensity increased with the magnitude of the sucrose concentration contrast: The pulsatile stimulus with the highest concentration difference (average sucrose concentration: 60 g/L) was rated as the sweetest in spite of the fact that the gross sucrose concentrations were identical over stimuli. Moreover, this stimulus was rated equally sweet as a continuous reference of 70 g/L sucrose. On alternation of sucrose with the qualitatively different citric acid, sweet taste enhancement remained at the level observed for alternation with water at citric acid concentration levels up to 3 times its detection threshold. Alternation of a sucrose solution with a citric acid solution at 9 times its threshold concentration, resulted in an attenuation of the pulsation-induced enhancement effect. Upon alternation of citric acid pulses at concentrations around the threshold with water intervals only, no taste enhancement was observed compared with continuous citric acid stimuli of the same net concentration. We propose that the magnitude of pulsation-induced taste enhancement is determined by the absolute rather than relative change of tastant concentration. This explains why 1) pulsation-induced sweet taste enhancement is determined by the magnitude of the sucrose pulse-interval contrast and 2) the alteration of citric acid with water does not enhance taste intensity at detection threshold level.     

The Hydrostatic and Hydrodynamic Volumes of Polyols in Aqueous Solutions and Their Sweet Taste

The tastes and solution properties of sugar alcohols were studiedin an attempt to illuminate the mechanism of sweet taste chemoreception.The SMURF method was used to measure taste time-intensity ofaqueous solutions of sugar alcohols and the results were interpretedusing the Stevens power function and kinetic parameters. Theapparent molar volumes, apparent specific volumes, partial molarvolumes, partial specific volumes and intrinsic viscositiesof the solutions were studied. Apparent molar volume reflectsthe size of the molecule in a hydrostatic state whereas intrinsicviscosity gives a measure of the size of the molecules in ahydrodynamic state. Generally the apparent molar volumes ofthe polyols are 6–13% greater than those of the parentsugars, indicating less interaction with the water structure.Apparent specific volume values can predict taste quality, andthe average apparent specific volume for the sugar alcoholsstudied fits within the central part of the sweet range, i.e.0.5–0.68 cm³/g, which accords with their ability to elicita pure sweet taste response. Intensities and persistences ofsweetness in the polyols followed the same trend as intrinsicviscosities. Chem. Senses 22: 149–161, 1997.     

Trying to detect taste in a tasteless solution: modulation of early gustatory cortex by attention to taste

Selective attention is thought to be associated with enhanced processing in modality-specific cortex. We used functional magnetic resonance imaging to evaluate brain response during a taste detection task. We demonstrate that trying to detect the presence of taste in a tasteless solution results in enhanced activity in insula and overlying operculum. The same task does not recruit orbitofrontal cortex (OFC). Instead, the OFC responds preferentially during receipt of an unpredicted taste stimulus. These findings demonstrate functional specialization of taste cortex in which the insula and the overlying operculum are recruited during taste detection and selective attention to taste, and the OFC is recruited during receipt of an unpredicted taste stimulus.     

Contribution of alpha-gustducin to taste-guided licking responses of mice

We examined the necessity of alpha-gustducin, a G protein alpha-subunit expressed in taste cells, to taste-mediated licking responses of mice to sapid stimuli. To this end, we measured licking responses of alpha-gustducin knock-out (Gus-/-) mice and heterozygotic littermate controls (Gus+/-) to a variety of 'bitter', 'umami', 'sweet', 'salty' and 'sour' taste stimuli. All previous studies of how Gus-/- mice ingest taste stimuli have used long-term (i.e. 48 h) preference tests, which may be confounded by post-ingestive and/or experiential effects of the taste stimuli. We minimized these confounds by using a brief-access taste test, which quantifies immediate lick responses to extremely small volumes of sapid solutions. We found that deleting alpha-gustducin (i) dramatically reduced the aversiveness of a diverse range of 'bitter' taste stimuli; (ii) moderately decreased appetitive licking to low and intermediate concentrations of an 'umami' taste stimulus (monosodium glutamate in the presence of 100 microM amiloride), but virtually eliminated the normal aversion to high concentrations of the same taste stimulus; (iii) slightly decreased appetitive licking to 'sweet' taste stimuli; and (iv) modestly reduced the aversiveness of high, but not low or intermediate, concentrations of NaCl. There was no significant effect of deleting alpha-gustducin on licking responses to NH4Cl or HCl.     

Taste confusions following gymnemic acid rinse.

The effect of a gymnemic acid (GA) rinse, which simulated a sweet-taste deficit, was measured on human taste perception and identification. Taste ratings showed that GA reduced the intensities of sucrose and aspartame to 14% of pre-rinse levels; over the recovery interval of 30 min, these values increased linearly to 63% of the pre-rinse levels. Repeated presentations of a set of 10 stimuli (five primarily or partly sweet--sucrose, aspartame, and NaCl-sucrose, acid-sucrose and quinine-sucrose mixtures; and five nonsweet--NaCl, KCl, Na glutamate (MSG), quinine HCl and citric acid) for identification following water and GA rinses produced 'taste confusion matrices' (TCMs). Correct identification of the sweet-tasting stimuli was reduced by 23% in presentations closely following the GA rinse, an effect that dissipated with time. Most misidentifications involved sucrose and mixtures containing sucrose. In a second TCM experiment, GA was presented frequently within each session to maintain the sweet taste deficit, which revealed itself as specific confusions. Rinsing with GA impaired discriminability of sweet-nonsweet pairs of stimuli but enhanced discriminability of the aspartame-(NaCl-sucrose) pair. GA had no effect on discriminability of nonsweet stimulus pairs. The results suggest that specific error patterns in the TCM could be used to identify quality-specific taste disorders.     

Amiloride is an ineffective conditioned stimulus in taste aversion learning in C57BL/6J and DBA/2J mice

The epithelial sodium channel (ENaC) blocker amiloride has been shown to increase the behaviorally measured NaCl detection threshold in mice. In this study, a conditioned taste aversion (CTA) paradigm was used to examine whether 100 microM amiloride has a perceptible taste that could contribute to this observed decrease in behavioral responsiveness. Eighty-four C57BL/6J (B6) and 64 DBA/2J (D2) mice were divided into eight groups (n=8-12 per group), in which half received an injection of 0.15 M LiCl (2 mEq/kg) and the other half an equivalent saline injection, in three conditioning trials. The four conditioned stimuli were 100 microM amiloride hydrochloride, water, 0.1 and 0.3 M NaCl. Neither strain demonstrated acquisition of a CTA to amiloride in a brief-access (BA) taste test (5 s trials in the gustometer). Although 0.3 M NaCl is inherently aversive, its pairing with LiCl led to significantly further decreases in licking during the BA test on salt trials in both strains. The D2 strain clearly avoided 0.1 M NaCl, whereas avoidance of this stimulus was more equivocal in B6 mice. The inefficacy of amiloride to serve as a conditioned stimulus in taste aversion learning involving three LiCl pairings suggests that the effects of this ENaC blocker on taste-related behavioral responses to NaCl are likely due to its pharmacological interference with sodium taste transduction.     

Taste performance on the anterior human tongue varles with fungiform taste bud density

This study demonstrated that individual diversities of tastesensitivity on the anterior tongue are due, in part, to variationsin fungiform taste bud density. Citric acid solutions were deliveredto a closed, spatially-matched (each test site was 43 mm²) flowchamber attached to the surface of the anterior tongue in 84subjects. A two-alternative forced choice, modified staircaseprocedure was used to derive a detection threshold value forcitric acid. The same session also included a visual analograting procedure to scale the taste intensity judgement of fiveconcentrations of citric acid. The taste buds within the chamberwere distinguished by methylene blue stain and recorded by videomicroscopy.The sip-and-spit method was used to contrast the spatially-matchedcondition with whole mouth stimulation. We found that detectionthreshold values were inversely related to the number of fungiformtaste buds, independent of gender or age. Whole-mouth thresholdvalues determined for each subject were always well below thespatially-matched threshold values. In addition, the data demonstratedthat subjects were able to scale the dynamic range of citricacid solutions. However, y-intercept approached zero intensityas the number of fungiform papillae decreased. The observeddifferences in citric acid sensitivity and fungiform taste buddensity indicated that taste performance on the anterior tonguein humans varies, in part, with the number of taste buds.     

SAMPLE/RINSE TEMPERATURE EFFECTS ON DISCRIMINATION OF SWEET AND SALTY DIFFERENCES IN A FOOD SYSTEM

The ability of panelists to distinguish between samples of apple juice with two levels of added sucrose (0.1 and 0.9 g/100 mL) or NaCl (0.05 and 0.14 g/100 mL) was determined using a modified signal detection technique. Samples at various temperatures (6, 24, or 50C) were presented with various rinse conditions. Sometimes a water rinse (6, 24, or 50C) was presented between samples and somtimes not. R-index values were calculated to predict the ability of panelists to distinguish between samples. No temperature effect was found for sweetness difference tests; however, R-indices showed that salty differences in apple juice were more distinguishable at a sample temperature of 24C than at 6 or 50C. Samples temperature had a greater effect than rinse condition (temperature or no rinse) on discrimination between samples differing in NaCl concentration.     

Responses of the rat chorda tympani nerve to glutamate-sucrose mixtures

Monosodium glutamate (MSG) has a multifaceted, unusual taste to humans. Rats and other rodents also detect a complex taste to MSG. Responses of the chorda tympani nerve (CT) to glutamate applied to the front of the tongue were recorded in 13 anesthetized rats. Whole-nerve responses to 30 mM, 100 mM and 300 mM MSG mixed with 300 mM sucrose were recorded before and after adding 30 micro M amiloride to the rinse and stimulus solutions. Responses of CT single fibers were also recorded. Predictions from models of whole-nerve responses to binary mixtures were compared to the observed data. Results indicated that MSG-elicited CT responses have multiple sources, even in an amiloride-inhibited environment in rats. Those sources include responses of sucrose-sensitive CT neural units, which may provide the substrate for a sucrose-glutamate perceptual similarity, and responses of sucrose-insensitive CT neural units, which may respond synergistically to MSG-sucrose mixtures.     

The influence of temperature on the threshold values of primary tastes

The threshold values of taste substances are influenced by severalfactors. To learn about the effect of the temperature on stimulus,recognition and terminal thresholds, these threshold valueswere determined for sucrose, sodium chloride, caffeine, quininehydrochloride, citric and tartaric acid at temperatures of 10,20, 40 and 60°C in a panel of 19 tasters. The individualvalues were found to vary over a wide range, resulting in arelatively large standard deviation of the mean threshold values.A temperature-dependence was found for the stimulus and recognitionthresholds which was different for the different taste substances.The stimulus and recognition thresholds are lowest in the temperaturerange of 20 to 40°C. The threshold values increase withincreasing temperature, and except for citric acid, significantand highly significant differences existed particularly between20 and 60°C, whereas statistically verifiable results couldnot be obtained between 10 and 20°C. There was no verifiabletemperature dependence either for the terminal thresholds. Theterminal thresholds were found to lie rather in a relativelynarrow concentration range and to be largely independent ofthe temperature. The results suggest that in warm dishes andbeverage more taste substances are required to produce the sametaste intensity. A dependence of the individual thresholds uponage, sex and smoking habits could not be found.     

Stimulus-response method for flows and volumes in slightly perturbed constant parameter systems

The area and the first time moment of tracer kinetic data on steady state systems contains information on flows and volumes of distribution in the system. A simple and general method of obtaining such information is given. The method consists of equating the ratio of the time integrals of appropriate stimulus function and response function in an actually performed finite amount tracer experiment to the ratio of stimulus and response that would exist in the tracer steady state. The generality of the method is illustrated by deriving the volume of osmotically exchangeable water in an organ from measurement of its transient bulk water exchange during a small osmotic perturbation.     

Oral chemical irritation: does it reduce perceived taste intensity?

In an experiment in which capsaicin was presented in physicalmixture with taste stimuli, no reduction in perceived tasteintensity, relative to the control condition, was observed.This result is inconsistent with previous reports of taste intensityreductions when oral chemical irritants were only periodicallyinterspersed among taste stimuli (Lawless et al., 1985; Lawlessand Stevens, 1984). A second experiment directly compared thetwo presentation formats and their respective control conditionsin a repeated-measures design. The results of that test confirmthat greater apparent reductions in taste intensity are observedwhen the oral irritant is presented as a periodic rinse eventhough perceived irritation under rinse conditions regularlyfalls to levels significantly below those maintained with mixturepresentations. This observation indicates that much of the apparentmasking of taste intensity in the presence of oral irritationis not directly related to irritation level but is sensitiveto procedural variation.     

THE EFFECT OF SOME BEVERAGES ON THE PERCEPTION OF SUCCEEDING BASIC TASTES

The objective of this study was to investigate the effect beverages have on subsequently perceived basic tastes. Two types of milk, Cola and water were used to rinse the mouth prior to conducting a Time Intensity study with basic taste solutions. From this some effects of the prior beverage onto the TI parameters was found. Especially Cola showed an effect on the intensity of the subsequently tasted sweet taste, and whole milk showed a stronger effect than skim milk. The effects showed both in size and timing of the basic taste intensities.     

A method for separate quantitative determination of 2-keto-3-deoxy-octonate and 3,6-dideoxyhexoses in mixtures.

A method for analysing ³²P in small aqueous samples by measuring the ?erenkov radiation is described. Centering problems with small sample volumes are eliminated by placing the sample in a polystyrene tube in the scintillation vial. The detection efficiency is high, 54.5 ± 0.6% (2 SD) at a sample volume of 25 μl. The reproducibility is good and independent of the sample volume. The detection efficiency of ³²P in polyacrylamide gel is shown to be as good as in water.     

Obese Women Have Lower Monosodium Glutamate Taste Sensitivity and Prefer Higher Concentrations Than Do Normal‐weight Women

The goal of this study was to determine whether obese women exhibit altered umami and sweet taste perception compared to normal‐weight women. A total of 57 subjects (23 obese and 34 normal weight) participated in a 2‐day study separated by 1 week. Half of the women in each group were evaluated using monosodium glutamate (MSG; prototypical umami stimulus) on the first test day and sucrose on the second test day; the order was reversed for the remaining women. We used two‐alternative forced‐choice staircase procedures to measure taste detection thresholds, forced‐choice tracking technique to measure preferences, the general Labeled Magnitude Scale (gLMS) to measure perceived intensity of suprathreshold concentrations, and a triangle test to measure discrimination between 29 mmol/l MSG and 29 mmol/l NaCl. Obese women required higher MSG concentrations to detect a taste and preferred significantly higher MSG concentrations in a soup‐like vehicle. However, their perception of MSG at suprathreshold concentrations, their ability to discriminate MSG from salt, and their preference for sucrose were similar to that observed in normal‐weight women. Regardless of their body weight category, 28% of the women did not discriminate 29 mmol/l MSG from 29 mmol/l NaCl (nondiscriminators). Surprisingly, we found that, relative to discriminators, nondiscriminators perceived less savoriness when tasting suprathreshold MSG concentrations and less sweetness from suprathreshold sucrose concentrations but had similar MSG and sucrose detection thresholds. Taken together, these data suggest that body weight is related to some components of umami taste and that different mechanisms are involved in the perception of threshold and suprathreshold MSG concentrations.     

Energy model for contrast detection: spatiotemporal characteristics of threshold vision

A model for contrast detection of spatiotemporal stimuli is proposed which consists of a spatiotemporal linear filter, an energy device and a threshold device. Assuming the existence of independent intrinsic noise, the probability of stimulus detection was approximated by a Weibull function of the response energy. With this assumption, the stimulus energy is a constant at fixed detection probability. This energy model for contrast detection satisfactorily accounted for the elliptical threshold contours of line pairs at stimulus separations within the range 2–30 min and at stimulus onset asynchronies within the range 20–140 ms. The threshold contour at a large stimulus onset asynchrony (300 ms) was in the form of a rounded square. This finding was explained by assuming that the probability of seeing the line pair was determined by the joint probability that at least one stimulus had been detected. With the energy model, the temporal and spatial autocorrelation functions of the response to a flashed line were evaluated. The autocorrelation functions thus determined were used to predict the temporal contrast sensitivity function to a flickering line stimulus and the spatial contrast sensitivity function to flashed gratings, which were in agreement with the experimental data. The data obtained were fitted adequately by an impulse response approximated by a spatiotemporal Gabor-like function. Received: 08 December 1997 / Accepted in revised form: 26 January 1999     

Adaptation to warming but not cooling at slow rates of stimulus change in thermal threshold measurements

The relationship between thermal detection threshold and rate of temperature change of the thermal stimulus when slow (<1 degrees C s(-1)) rates of change are employed was investigated. Using both the reaction time (RT) inclusive Method of Limits and RT exclusive Method of Levels healthy volunteers had warming (WDT) and cooling detection thresholds (CDT) measured at four different rates of temperature change (0.3, 0.5, 0.7 and 1.0 degrees C s(-1)) from the thenar and/or mental regions using a contact thermode. With the Method of Limits, CDT increased linearly with rate of temperature change suggesting increments were due to RT artefacts. This was further supported by threshold assessment with the Method of Levels which showed CDT were unaffected by the rate of change in the RT exclusive method (P > 0.1). In contrast, WDT did not increase linearly with rate of stimulus temperature change when the Method of Limits was used and threshold assessment with the Method of Levels showed WDT assessed using a 0.3 degrees C s(-1) ramp rate were significantly higher than those measured with a 1 degrees C s(-1) rate of change (P < 0.05). This study indicates that adaptation to a warming stimulus can occur at faster rates of stimulus change than previously anticipated and identifies differences in warming and cooling pathways in sensitivity to adaptation.     

A brief-access test for bitter taste in mice

Inbred mouse strains vary in their response to bitter-tasting compounds as assessed by 48 h preference tests. These differences are generally assumed to result from altered gustatory function, although such long-term tests could easily reflect additional factors. We developed a brief-access taste test and tested the responses of two inbred strains, as well as C3. SW congenic mice, to the bitter stimulus sucrose octaacetate (SOA). Water-deprived trained mice were tested with five concentrations of SOA (0.00018-0.18 mM) and distilled water in a Davis MS- 160 apparatus. Trials were 5 s in duration and stimuli were presented randomly within blocks; each stimulus trial was preceded by a water rinse trial. Each concentration was presented twice in a session and mice were repeatedly tested across consecutive days. SOA-taster mice, including the SWR/J (SW) inbred and C3. SW congenic taster (T) mice, avoided licking SOA at concentrations >0.003 mM. In comparison, C3HeB/FeJ (C3) and C3. SW demitaster mice (D) licked all concentrations at the same rate as water. Concentration-response functions were similar across strains for both the brief-access test and a parallel 48 h preference test run on separate groups of mice. Furthermore, concentration-response functions were similar whether or not the brief-access test was preceded by a 4 day, single concentration pretest with SOA. The brief-access test is a suitable assay for bitter taste function in mice because it minimizes possible post-ingestive influences on taste.