Chem. Senses 27: 31-38,
2002
© Oxford University Press 2002
Bitter Taste of Saccharin and Acesulfame-K
Department of Food Science, Cornell University, Ithaca, NY 14853
Correspondence to be sent: Harry Lawless, Department of Food Science, Cornell University, Ithaca, NY 14853, USA. E-mail: htl1{at}cornell.edu
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Abstract |
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 Top Abstract IntroductionExperiment 1: correlations among...Experiment 2: re-examination...DiscussionReferences |
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The relationships among suprathreshold taste responses to acesulfame-K, Na-saccharin and 6-n-propylthiouracil (PROP) were examined in two studies. In the first study, the labeled magnitude scale was used with the high anchor labeled as `strongest imaginable oral sensation' and in the second study, it was labeled as `strongest imaginable sensation of any kind'. Results from the two procedures were similar. Individual differences among 65 subjects were seen in bitter responses to acesulfame-K and saccharin. Bitter responses to acesulfame-K ands accharin were positively correlated, but showed no significant relationship with responses to PROP bitterness or with PROP taster groups. Saccharin and acesulfame-K may share a common mechanism for bitter taste reception and transduction, one that varies across individuals and is different from mechanisms mediating bitter responses to PROP. Changing the instructions of the labeled magnitude scale induced a context effect. Ratings of sweetness referenced to the `strongest imaginable sensationof any kind' were lower than ratings referenced to just oral sensations.
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Introduction |
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TopAbstractIntroductionExperiment 1: correlations among...Experiment 2: re-examination...DiscussionReferences |
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Salts of saccharin are intensely sweet but also have a bitter taste to some individuals. Helgren et al. (Helgren et al., 1955
) estimated that 
25% of a European population will detect
an off-taste to saccharin described as metallic or bitter. For a time, it was
proposed that this unpleasant nonsweet taste was due to impurities in the
stimulus, specifically o-toluene sulfonamide, a by-product produced
during synthesis (McKie, 1921;
Cannon, 1954). However, other
research showed that the off-tastes were a true property of purified saccharin
salts (Rader et al.,
1967). Using time-based psychophysical methods, Larson-Powers and
Pangborn (Larson-Powers and Pangborn,
1978) showed saccharin to have a lingering bitterness. The
bitterness is more pronounced at higher concentrations, producing a flattening
of the sweetness function (Moskowitz and
Klarman, 1975) and making iso-sweet matches to other compounds
difficult to achieve at high levels (Ayya
and Lawless, 1992). Acesulfame-K is another intense sweetener with
a bitter taste. According to Ott et al.
(Ott et al., 1991),
acesulfame-K has a delayed bitter aftertaste that is more intense and longer
in duration than sucrose, aspartame or alitame. Schiffman et al.
(Schiffman et al.,
1985) noted high variability in the intensity and quality of
acesulfame-K, due to bitter and metallic sidetastes.
Bartoshuk (Bartoshuk, 1979)
found a relationship between 6-n-propylthiouracil (PROP) taster
status and saccharin bitterness, with PROP tasters being more sensitive to
sodium saccharin bitterness at low levels (0.001 and 0.0032 M). At higher
concentrations, the group differences attenuated. Since that report, a third
group of people with very high PROP sensitivity has been classified as
supertasters. These individuals have a pronounced response to suprathreshold
concentrations of PROP when assessed relative to NaCl standards. They also are
characterized by having a higher density of taste buds and fungiform papillae
and lower PROP thresholds than tasters and non-tasters
(Reedy et al., 1993;
Bartoshuk et al.,
1994; Drewnowski et
al., 1997). Drewnowski et al.
(Drewnowski et al.,
1997) found a higher level of rated bitterness for 0.0032 M
Na-saccharin among tasters and supertasters of PROP than among non-tasters,
but no difference among the groups at higher levels. Another study
(Gent and Bartoshuk, 1983)
collected bitterness responses from PROP tasters and non-tasters but found no
differences in ratings when solutions were flowed across the front of the
tongue. Nor did they not report any differences for solutions that were sipped
in a whole-mouth stimulation method. Ly and Drewnowski
(Ly and Drewnowski, 2001)
noted that PROP taster differences in the perception of caffeine bitterness
were attenuated in taste mixtures.
Informal observations of individual differences in response to acesulfame-K
bitterness at higher concentrations raised the following question. Are
individuals who are sensitive to the bitter taste of saccharin also sensitive
to the bitter taste of acesulfame-K? There are some similarities in the
chemical structures, such as the negatively charged nitrogen in the ring
structure adjacent to an SO2 group and acarbonyl (see
Figure 1). These structural
similarities are consistent with the possibility of common stimulation and
transduction mechanisms. A number of different mechanisms have been proposed
for bitter taste transduction (Spielman
et al., 1992). Given the observation of wide individual
differences in the perception of saccharin and acesulfame-K bitterness,
transduction mechanisms for bitter taste other than those responsible for PROP
tasting might contribute to these individual differences. Thus we examined the
degree of correlation among PROP, saccharin and acesulfame-K tastes across
individuals.
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After we had conducted the first experiment, Bartoshuk
(Bartoshuk, 2000) suggested
that the specific instructions to subjects using labeled magnitude scaling
(LMS) (Green et al.,
1993,
1996) could obscure group
differences among PROP tasting groups due to personal context effects.
Specifically, supertasters, who live in a world of extreme taste sensations,
might show attenuated ratings of the stimuli presented during an experiment.
In other words, they might rate the moderately intense experimental stimuli
lower than expected due to their implicit comparison to the more intense oral
sensations they commonly experience. Contrast effects are common in human
judgements (Lawless et al.,
2000) and could obscure higher subjective intensities actually
experienced by this group within an experimental session. Bartoshuk suggested
that instructions should encourage subjects to use the scale with reference to
all sensations experienced in life (not just oral sensations) so that a more
equivalent contextual basis for scale usage could be achieved. Put more
simply, such instructions would provide a common and comparable frame of
reference for the taster groups. In accordance with this suggestion, we
conducted a second experiment using modified instructions for the LMS without
reference to oral sensations but rather instructing subjects to consider the
high end as the `strongest imaginable sensation of any kind' (italics
added).
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Experiment 1: correlations among PROP, saccharin and acesulfame-K |
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TopAbstractIntroductionExperiment 1: correlations among...Experiment 2: re-examination...DiscussionReferences |
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Methods and procedures
Subjects
Thirty-eight volunteers were recruited from the Cornell community (23
females, age range 20-55 years). Twenty-one of these individuals had
previously participated in a similar study [experiment 1 of
(Sposato, 2000)], and were
familiar with the rating scales and techniques employed. Subjects were paid a
token incentive for participation and signed an informed consent form
explaining the risks and voluntary nature of the procedures.
Stimuli
Standards for the training session consisted of 0.009 mM quinine
hydrochloride (form. wt 360.9) as the bitter standard and 4.8 mM citric acid
(form. wt 210.1) as the sour standard. Stimuli in the test sessions consisted
of the following substances and concentrations: NaCl (form. wt 60.0) at 0.32
and 1.0 M, Na-saccharin (form. wt 202.2) at 0.4 and 2.1 mM, acesulfame-K
(form. wt 201.2) at 1.2 and 5.2 mM and PROP (form. wt 170.2) at 1.0 and 3.2
mM. Sweetener concentrations were chosen on the basis of equations from Dubois
et al. (Dubois et al.,
1991) to produce sensations approximately equal to 100 and 225 mM
sucrose, representing weak-to-moderate levels of sweet taste intensity.
Concentrations of NaCl and PROP were chosen based on the screening tests used
by Bartoshuk et al. (Bartoshuk
et al., 1994). Spring water (Chemung Spring Water,
Chemung, NY) served as the diluent and rinse. Sample volumes were 30 ml each
and sample temperature was 20 ± 2°C. Samples were labeled with
random three-digit codes, using different codes for samples that appeared in
multiple sessions.
Procedures
Subjects participated in three experimental sessions. In the first session
subjects were screened for their abilities to categorize sour and bitter
sensations. Sensations of sourness and bitterness were described to the
subjects using examples of lemons for sourness and dark beer, strong coffee,
coffee grounds and tonic water for bitterness. Subjects were next presented
with a set of three samples, two containing the sour standard and one
containing the bitter standard, or vice versa. Samples were coded with random
three-digit codes and presented in random order. Subjects tasted the samples
and categorized them as sour or as bitter. All subjects correctly categorized
their three samples.
In the second and third sessions, subjects rated each of the eight test samples, two concentrations each of PROP, NaCl, saccharin and acesulfame-K. Sweeteners and NaCl were presented first in random order. PROP samples were tasted last in order of increasing concentration. This was done to prevent a strong persistent bitter sensation from affecting the evaluation of other samples in the case of PROP-sensitive individuals. Subjects took the entire sample into their mouths, circulated it for 15 s to cover all oral surfaces and expectorated. Immediately upon expectoration, subjects rated taste intensity for the attributes of sweetness, saltiness, sourness and bitterness using a horizontal version of the LMS with the upper bound of the scale labeled as `strongest imaginable oral sensation.' Subjects rinsed their mouths with spring water between each sample. Responses were made by placing a mark on the LMS line displayed on a computer screen, using a mouse. Data were collected using the Compusense 5 System (Compusense, Inc., Guelph, Ontario, Canada).
Analysis
Data were exported from the Compusense system for statistical analyses
using Statistica (v.5.1, Statsoft, Tulsa, OK). Analyses included repeated
measures ANOVA (subjects random, with PROP taster status as a between-groups
factor), simple correlations and principal components analysis (PCA). Varimax
normalized rotated solutions were found and factors retained that explained
>10% of the total variance. Replicates from the two sessions were averaged
after finding that none of the bitter ratings showed a significant replicated
difference in ANOVA.
Results
Subjects were classified into PROP non-tasters, tasters or supertasters,
based on their PROP ratio (Bartoshuk et
al., 1994), defined as follows:
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Subjects with PROP ratios of <0.4 were classified as non-tasters (n = 11), between 0.4 and 1.2 as tasters (n = 14) and >1.2 as supertasters (n = 13).
Figure 2 shows the sweetness and bitterness of each sweet compound. A significant concentration effect was found for both substances for all taste qualities [all F(1, 73) > 7.6, P < 0.001]. As shown in Figure 3, the bitterness responses for the taster groups were distributed throughout the scale range. There were no group differences nor interactions with PROP taster groupings.
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Correlations showed the expected positive relationship between bitterness ratings for 1 and 3.2 mM PROP (r = +0.86, P < 0.001). A positive correlation was also seen among the pairs of saccharin and acesulfame-K stimuli, as shown in Table 1. The relationship was especially strong at the higher levels of the sweeteners, where bitterness is more apparent to some individuals. The correlation of +0.84 (see Figure 4) was almost as high as the correlation between the two levels of PROP bitterness (probably a ceiling due to measurement error) and shows a strongly related pattern of individual differences.
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However, correlations between acesulfame-K bitterness and PROP bitterness were not significantly different from zero, and the same was true for saccharin bitterness and PROP bitterness. No significant correlations were observed between PROP bitterness ratings and sweetness ratings of either sweetener.
PCA also showed this pattern of correlation. The varimax normalized loadings for the first six factors (70% explained variance) are shown in Table 2. The first factor subsumed the largest amount of variance (22%) and may reasonably be interpreted as a general intensity dimension that correlated with many of the side tastes of the stimuli. Saccharin and acesulfame-K sweetness ratings loaded on the second factor and their bitterness loaded on factor 6, separately from the PROP bitterness ratings and the PROP ratio index, which loaded on factor 3. Other rotation options (varimax unnormalized, equamax, quartimax) and unrotated solutions were also examined, and in all cases PROP and bitterness of the sweeteners were loaded on separate factors.
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Experiment 2: re-examination with modified LMS anchors |
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TopAbstractIntroductionExperiment 1: correlations among...Experiment 2: re-examination...DiscussionReferences |
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Methods and procedures
Subjects
Thirty volunteers from the Cornell community (16 female) completed the
study. None of these subjects had participated in experiment 1. Subjects were
selected for the study in order to produce equal numbers ofPROP non-tasters,
tasters and supertasters (10 per group, classified by the same criteria as in
experiment 1), and to have approximately equal gender balance in each group.
Seven additional subjects failed to correctly categorize the bitter and sour
stimuli in the screening test, so were dismissed from further participation.
Subjects were paid a token incentive for participation and signed an informed
consent form explaining the risks and voluntary nature of the procedures.
Following the third session, one supertaster and one non-taster were dropped
from the analysis for rating PROP as sweet, and one taster for rating PROP as
sour (sour/bitter confusion), leaving nine subjects per PROP group.
Stimuli
Samples were the same as in experiment 1 except that deionized water served
as the diluent and rinse water.
Procedures
The screening task for sour/bitter categorization was modified to include
three quinine samples and three citric acid samples. Panelists were required
to classify all six correctly into sour and bitter groups. ROP and NaCl
stimuli were rated in the second and fourth sessions, and the sweetener
samples were rated in duplicate in the third session. Ratings on all scales
were modified to change the upper bound label to `strongest imaginable
sensation of any kind'. Statistical analyses were the same as in experiment
1.
Results
Changing the anchoring instructions to the most intense experience rather than most intense oral sensation induced a contextual shift (Figure 5). Ratings of sweetness were 32% lower in this experiment than in experiment 1 [F(1,124) = 18.63, P < 0.001]. Bitterness ratings were also lower by 24% [F(1,124) = 3.00, P < 0.10]. The shift in ratings was similar for all three PROP taster groups (no group effect or interaction).
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Figure 6 shows the mean ratings of the two sweeteners for the three groups. For bitterness, there were significant interactions of PROP status with sweetener [F(2,51) = 4.23, P < 0.05] and concentration factors [F(2,51) = 9.77, P < 0.01], as well as a concentration by substance interaction [F(1,51) = 5.63, P < 0.05]. As shown in Figures 6 and 7, the PROP interactions were in an unexpected direction, with some PROP non-tasters giving higher bitterness ratings than the other groups to 5.2 mM acesulfame-K and slightly higher ratings to 2.1 mM saccharin. Supertasters gave higher sweetness ratings to the higher level of saccharin than did non-tasters (LSD test, P = 0.056). No other group differences were observed on the basis of PROP status.
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Examining the correlation pattern, once again bitterness ratings for the two PROP stimuli were correlated as expected (r = +0.69, P < 0.001). Significant correlations were found for bitterness ratings between the higher levels of acesulfame-K and saccharin (r = +0.64, P < 0.001), between the lower levels of the two sweeteners (r = +0.38, P = 0.051) and between the bitterness ratings of 2.1 mM saccharin and 1.2 mM acesulfame-K (r = +0.51, P < 0.01). One significant but negative correlation was observed between the PROP ratio and 5.2 mM acesulfame-K bitterness (r = -0.40, P < 0.05). In the correlation pattern there was evidence of a weak but positive relationship of PROP responses to sweetness intensity. The bitterness of 3.2 mM PROP was correlated with the sweetness of 5.2 mM acesulfame-K (r = +0.41, P < 0.05), with 1.2 mM acesulfame-K (r = +0.38, P < 0.05) and with 2.1 mM saccharin (r = +0.34, P < 0.10).
Factor loadings following the PCA are shown in Table 3. A general intensity factor was seen in the first factor, and PROP bitterness was included in factor 2. Factor 3 captured the variance associated with bitterness of the four saccharin and acesulfame-K stimuli, suggesting a pattern of inter-correlation orthogonal to the other ratings and separate from PROP. Bitterness indices were constructed in an analogy to the PROP ratio, taking the sum of bitterness ratings for each sweetener and dividing by the saltiness of NaCl. The saccharin index and acsulfame-K index were positively correlated (r = +0.47, P < 0.05), but neither was significantly correlated with the PROP ratio.
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Discussion |
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TopAbstractIntroductionExperiment 1: correlations among...Experiment 2: re-examination...DiscussionReferences |
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There are three different approaches to examination of the overlap or relationships among taste compounds. One is cross-adaptation, in which the adaptation to one compound is tested against various subsequent test stimuli (McBurney et al., 1972
). Reduction in the taste intensity of the second compound is
interpreted as evidence for common receptor or transduction mechanisms. This
provides the most direct psychophysical evidence for common pathways, but has
been so far limited to areas of the dorsal anterior tongue that can easily be
stimulated with flowing solutions. The second common approach is
preclassification of taster/non-taster groups (e.g. for PROP tasting), with
subsequent tests of group differences for sensitivity to a test compound or
comparison of suprathreshold responses
(Bartoshuk, 1979). This
approach requires a control for other differences among the classified groups,
such as a difference in general taste sensitivity or scale usage tendencies
(Delwiche et al.,
2000). A third approach is to examine correlations among
thresholds or among suprathreshold responses for pairs of test compounds
(Yokomukai et al.,
1993; Delwiche et
al., 2000). Unlike the `group differences' approach, this
allows an estimation of common variance that may reflect the degree of overlap
in taste receptor and transduction mechanisms for the two compounds. This
approach also provides a graded estimate of the overlap, rather than a binary
decision about a null hypothesis. The present experiments used both the group
comparison and correlational approaches to assess the commonalities among
bitter responses to PROP, saccharin and acesulfame-K.
These studies confirm the notion that there are individual differences in
responsiveness to suprathreshold bitterness of Na-saccharin and acesulfame-K,
and that these patterns of sensitivity are correlated across individuals. If a
person finds a strong bitter taste to a solution of Na-saccharin, he or she is
likely to find that an equi-intense solution of acesulfame-K is also strongly
bitter. Such responses, however, are not well predicted by PROP status. This
is consistent with the findings of Bartoshuk and Drewnowski et al.
(Bartoshuk, 1979;
Drewnowski et al.,
1997), who found a PROP taster difference for very low levels of
saccharin but no difference at moderate-to-high intensity levels. Ly and
Drewnowski (Ly and Drewnowski,
2001) also noted attenuated differences between PROP taster groups
for the bitterness of caffeine when a sweetener was added, and hedonic
differences between the groups were eliminated. Differences between PROP
taster groups and the patterns of correlation may be easier to uncover with
simple bitter stimuli than with more complex tastes, such as these
sweet—bitter stimuli, or with real foods.
Examining the bitter ratings of the sweet compounds, considerable residual variability persists beyond the shared variance of saccharin and acesulfame-K and beyond PROP status. In addition to some shared mechanisms of bitterness reception and transduction, there may be non-overlapping modes of stimulation, other chemical or peri-receptor access factors coming into play, or even general taste sensitivity differences between individuals that are unrelated to a saccharin bitterness dimorphism.
In spite of modification of the LMS verbal anchor, the patterns of results
from both studies were similar in terms of the bitterness correlations. As
noted in the introduction, one could predict paradoxically low responses from
PROP supertasters as a kind of context effect if their frame of reference is a
generally more intense set of oral experiences
(Bartoshuk, 2000). This is
reasonable given that the LMS, like other scaling methods, is prone to context
effects that influence the frame of reference in the subject's scaling
behavior (Lawless, et al.,
2000). The search for a scaling method that would provide a common
metric across individuals is still desired for comparison purposes, and
methods such as cross-modality matching and magnitude matching have been
applied for that purpose [for an overview, see Bartoshuk
(Bartoshuk, 2000)]. Borg's
assertion for the category-ratio scale was that having a comparison to the
strongest imaginable sensation might provide a common frame of reference
(Borg, 1982). Although this
seems reasonable, there is no direct evidence to support this assertion.
A general contextual shift in the form of contrast occurred, wherein the
more intense imagined experience led to lower intensity ratings for all taster
groups. The most intense experience of any kind for most people would
presumably be more intense in memory than the most intense oral experience.
Judgements in the context of overall experience were lower than judgements in
the context of oral experience. This effect is consistent with the findings of
Green et al. (Green et
al., 1996), who noted a lowered response range when LMS
ratings were cross-referenced to oral sensations rather than specific taste
qualities. One would expect oral sensations that include sensations like chili
pepper to be more intense than sensations of sweetness. Relative to this more
intense frame of reference, ratings are lowered, a form of contrast.
Given the diversity in bitter molecules and the genetic diversity in both
humans and other mammals in their sensitivities to different bitter compounds,
bitter taste transduction is likely to involve multiple mechanisms (Speilman
et al., 1996). One important mechanism appears to be blockage of
outward potassium flow by substances such as the potent bitter compound
denatonium (Brand, 1997).
Denatonium also appears to result in a release of intracellular calcium
stores, possibly mediated through a second messenger system. IP3 and diacyl
glycerol (DAG) have both been implicated as second messengers in bitter taste
transduction. DAG controls a protein kinase and IP3 could affect the release
of calcium from intracellular stores. Another possibility is that the
activation of gustducin in a G-protein-coupled receptor system stimulates a
phosphodiesterase enzyme. This would somewhat paradoxically cause a decrease
in cyclic nucleotides, such as cAMP. If cAMP is gating an inhibitory channel,
lower levels could stimulate membrane depolarizing events
(Brand, 1997). Recently, a
novel family of 40-80 G-protein-coupled receptors have been identified in
humans and mice that respond to bitter tastants
(Adler et al., 2000;
Chandrashekar et al.,
2000). More than one receptor is apparently expressed within the
same taste cell, which would help explain why chemically diverse bitter
tastants give rise to similar sensations. At this time, it is not known what
mechanisms of bitter taste transduction are activated by saccharin and
acesulfame-K. However, the present results support the notion of some overlap
of those mechanisms, mechanisms which differ among individuals.
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Acknowledgments |
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Supported by NIH grant RO1 DC-00902. Some of these data are from the MS thesis of Domenic Sposato, and were presented at the 2000 and 2001 meetings of the Association for Chemoreception Sciences. The order of authorship is alphabetical, reflecting equal contributions.
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Accepted September 10, 2001
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T. Ohkuri, K. Yasumatsu, N. Horio, M. Jyotaki, R. F. Margolskee, and Y. Ninomiya Multiple sweet receptors and transduction pathways revealed in knockout mice by temperature dependence and gurmarin sensitivity Am J Physiol Regulatory Integrative Comp Physiol, April 1, 2009; 296(4): R960 - R971. [Abstract] [Full Text] [PDF]
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C. E. Riera, H. Vogel, S. A. Simon, and J. l. Coutre Artificial sweeteners and salts producing a metallic taste sensation activate TRPV1 receptors Am J Physiol Regulatory Integrative Comp Physiol, August 1, 2007; 293(2): R626 - R634. [Abstract] [Full Text] [PDF]
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B. G. Green and P. George 'Thermal Taste' Predicts Higher Responsiveness to Chemical Taste and Flavor Chem Senses, September 1, 2004; 29(7): 617 - 628. [Abstract] [Full Text] [PDF]
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S. V. Kirkmeyer and B. J. Tepper Understanding Creaminess Perception of Dairy Products Using Free-Choice Profiling and Genetic Responsivity to 6-n-Propylthiouracil Chem Senses, July 1, 2003; 28(6): 527 - 536. [Abstract] [Full Text] [PDF]
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