Components of Flavour
I
Interactions among salty,
sour and bitter compounds
Paul AS. Breslin
The human gustatory system is capable of responding
processing the taste of solitary compounds in water. How-
ever, the taste system rarely contacts solitary compounds out-
side the laboratory and has surely evolved to process complex
mixtures of sapid chemicals, such as occur in virtually
foods. This review will focus primarily on the lesser-studied
interactions between pairs of salty, sour and bitter compounds.
Pair-wise interactions among these three taste qualities should
be of interest because they constitute a significant proportion
(-30-50%) of possible binary taste interactions. In general,
salts and acids enhance each other at moderate
trations but suppress each other at higher concentrations.
Bitter compounds and acids can either enhance or suppress
each other depending on the concentrations, the food stimuli
and the experimental methods involved. Sodium
bitter compounds generally interact so that bitterness
suppressed to some variable degree and the saltiness is un-
affected. As will be described below, there are exceptions to
all of these generalities.
When two compounds that elicit different taste qualities
are mixed in solution at moderate or strong concen-
trations, the mixture will often yield a taste sensation that
is less intense than the simple sum of the component
tastes. In two-component mixtures, each of the taste
qualities is usually suppressed (perceived as less intense)
than when it is tasted separately’-4 (see Box 1 and Fig. 1).
Binary combinations of certain taste stimuli may also
result in extremely asymmetrical suppressions. For ex-
ample, Schifferstein and Frijters”, and Breslin and
Beauchamp12 reported that when quinine hydrochloride
(QHCl), which usually elicits a bitter taste, was mixed
in solution with sodium chloride (NaCl), which usually
elicits a salty taste, the saltiness of the NaCl was rela-
tively unaffected, whereas the bitterness of the QHCl
was suppressed by 50-70% (see also Ref. 13), depend-
ing on the concentrations involved (contrast with Refs 3
and 14). Although taste suppression is commonly ob-
served, in no instance has a parallel been found in taste
to the color opponency phenomenon, in which two
lights cancel each other’s qualities15. In fact, in taste, the
Paul AS. Breslin is at the Monell Chemical Senses Center, 3500 Market
Street, Philadelphia, PA 19104, USA (fax: +l-215-898-2084;
breslin@cattell.psych.upenn.edu).
390 Copyrlght01996. Elsewer Science Ltd. All rights reserved. 0924 -2244/961$15.00
PII:50924-2244(96)10039-X
opposite is often true, as when two weak compounds
enhance one another2,‘J6.
Moskowitz” carefully explored whether the enhance-
ment or suppression seen in many mixtures was either
true synergism or true masking (see Box 1 and Fig. 3).
That is, he determined whether the shape of the con-
centration-intensity function (‘power function’; i.e. the
perceived intensity plotted as a function of the con-
centration of tastant) for a single compound changed
when tasted against a background taste. He found that
to and the power functions shifted laterally but the exponents
(slopes) of the functions did not change. In other words,
the rate of growth of the intensity as a function of con-
centration was unaltered by the presence of a back-
ground compound. A leftward shift of the power func-
all tion is defined as enhancement by the background taste.
A rightward shift is defined as suppression by the back-
ground taste. When the slope of the function changes to
become either steeper or shallower (a rare occurrence in
tastant mixtures), then the background compound is af-
fecting the rate of growth of taste intensity as a function
of concentration, which is defined as synergism or
concen- masking (see Box 1 and Fig. 3).
There are several reports of binary tastant mixtures
resulting in the enhancement of individual taste qualities.
Reliable enhancement has been reported with mixtures
such as combinations of sweetenersi0*i8.i9, mixtures of
salts and monosodium glutamate (MSG) with 5’-ribonucleotides9~*o,
is or mixtures of sucrose with dilute NaC121.Interestingly,
the enhancements seen in mixtures of 5’ribonucleotides
and MSG and in certain combinations of intense sweet-
eners are the only apparent cases of synergism that have
been identified, in which the taste of the two com-
ponents in mixture is greater than any linear model
would predict based on their tastes when unmixed9*‘0~‘9.
One of the more striking features of the literature on
heterogeneous taste mixtures is the lack of agreement
across laboratories. There are several reasons for this that
should be kept in mind when evaluating the literature:
(1) Differences in individual subjects’ sensitivities to vari-
ous taste compounds can be quite pronounced and this
results in very different responses to mixtures of the
compounds22,23.
(2) There is a tendency within the field to expect all
compounds that share a common taste quality (e.g. all
bitter-tasting compounds) to interact with other com-
pounds (e.g. all sour-tasting compounds) in a similar
way. This has resulted in questions such as ‘How do
sour stimuli interact with bitter stimuli?’ being asked.
This question cannot be answered simply, because even
if all sour stimuli elicit the same sourness quality and all
bitter stimuli elicit the same bitterness quality, they are
likely to do so through multiple transduction mecha-
nisms24-26;and enhancement and suppression may, in
the case of some interactions, involve transductive path-
ways directly27-29. Because the effects of some taste
mixtures will be mechanism specific, one cannot expect
e-mail: all bitter-tasting stimuli to behave the same way in the
presence of an acid (or acids) and vice versa.
Trends in Food Science & Technology December 1996 [Vol. 71
Special Issue on Flavour Perception
 Box 1. Definitions of types of potential interactions between two mixed co~~~pounds
linear
Suppression: (for suprathreshold concentrations)
Suppression is a linear (or subtractive) process that bears direct
analogy with subthreshold hypoadditivd mixing as well as with
subtractive mixing (see below), except that it is applied here to
suprathreshold compounds. As shown in Fig. 1, it is most clearly
I depicted by a rightward shift of the concentration-intensity func-
tioh along. the concentration axis in the presence of a second
compound, so that every concentration point along the curve is
per&&d as less intense. One should no& that the-function itself
is not finear and will not result in the same magnitude decrease
at different positions along the concentration-in&&y curve (see
Fig. 2P.
1Enhancement:(for suprathreshold concentrations)
Enhancement is a linear (or additive) process that bears direct anal-
ogy with subthreshold hyperadditive mixing (see below), except
that it is applied here to suprathreshold compounds. As shown in
Fig. I, it is most clearly depicted by a iefhrvard shift of the concen-
tration-intensity function along the concentration axis in the pres-
ence of a second compound, so that every concentration point
along the curve is perceived as more intense. Again, one should
note that the function itself is not linear and will not result in the
same magnitude increase at different positions along the concen-
tration-intensity curve (see Fig. 2)jr6,
Additivemixing:(for subthreshold concentrations)
Additive mixing may be the same process that governs enhance-
ment and suppression. Additive mixing involves the determination
of the threshold concentration for compound A in the presence of
a subthreshold concentration of compound B. Simple (completet
additivity means that, when the concentrations of A and B are re-
duced below their individual threshold concentrations and mixed
together, the detection threshold for the mixture will occur
when each compound is at 50% of its concentration threshold (or
30 : 70; 70 : 30; 40 : 60; 60:40, etc.), so that the sum of the per-
centages of the individual threshold concentrations for A and B
equals 100%. This usually results in nearly additive combinations
of compounds A and B (Ref. 7).
Hvwradditive mixine:(for subthreshold concentrationsl
Hypradditive mixin; is defined in the same way as a;fditive mix-
ing except that the threshold for the mixture is obtained when the
compounds are at concentrations corresponding to percentages of
(3) Several (perhaps all) strong suprathreshold taste mix-
tures display non-monotonic interactions (the direction
of the interaction changes, sometimes increasing taste
and sometimes decreasing taste, depending on the initial
magnitude of the taste in question). For example, NaCl
enhances the sweetness of sucrose when the NaCl is
present at a low concentration, has no effect when pres-
ent at a moderate concentration, and suppresses the
sweetness of sucrose when present at a high concen-
tration2’ (see ‘Conclusions’ section). Given that many
researchers work with one or only a few concentrations,
and different laboratories select different concentrations
with which to work, disagreement seems inevitable.
(4) Different experimental protocols will often yield dis-
crepant results. For example, whole-mouth sipping ex-
poses the stimuli to a much greater array of potential
taste receptor interactions (palatine and epiglottal as well
as pharyngeal) than would a simple lingual dorsal-flow
design; that is, compounds will taste differently when
-90% of taste buds are stimulated (as in ‘sip-and-spit’
Trends in Food Science & Technology December 1996 [Vol. 71
of dierent tastes
their individual thresholds that, when totalled, sum to less than
WW’
I”U 10 ‘I\
rthlj.
a 45% for A and 50% for B, or 45% for A and 45% for 8,
etc.)7. WY+~-~,~.
*or*Adi+i\re mixing bears a direct parallel to the supra-
A”,“t...r.l p~~~~~“lllcnon
A.‘.,,...,
u~~~~~~“Iu of enhancement.
Hypoaddiivemixing:(for subthreshold concentrations)
Hypoadditive mixing is defined in the same way as additive mixing
except that the threshold for the nmixture is obtained when the
compounds are at concentrations corresponding to percentages of
their individual thresholds that, when totalled, sum to more than
100% (e.g. 55% fur A and 50% for B, or 55% for A and 55% for B,
etc.17, Hypoadditive mixing bears aI direct parallel to the supra-
thr&hoId phenomenon of suppression.
Subtractivemixing:(for subthreshold and suprathreshold mixtures)
Subtractive mixing involves the determination of a threshold for
compound A in the presence of a suprathreshold background con-
centration of compound 6 (Ref. 7). This usually results in com-
pound B elevating the threshold for compound A.
Nonlinear
A&I&~: (for suprathreshold concentrations)
Masking is a nonlinear process in which the addition of compound
A decreases the intensity of compound B in a manner that goes
beyond iinear reductions in intensity. As shown in Fig. 3, it is most
clearly depicted by a decrease in the slope of the function and a
rightward shift of the concentration-intensity function along the
concentration axis in the presence of compound B, so that every
concentration point along the curve is perceived as less intense.
One should note that the function itself is not linear and will not
result in the same magnitude decrease at different positions along
the concentration-intensity curve (see Fig. 2P-“‘.
Synergism:[for suprathreshold concentrations)
Synergy is a nonlinear process in which the addition of compound
A increases the intensity of compound B in a manner that goes
beyond linear increments in intensity. As shown in Fig. 3, it is most
clearly depicted by an increase in the slope of the function and
a leftward shift of the concentration-intensity function along the
rnnrontrs*ion axis in the presence of compound B, so that every
C”II\.~Ilil~I
concentrat ion point along the curve is perceived as more intense.
One should note that the function itself is not linear and will not
result in the same magnitude increase at different positions along
the concentration-intensity curve (see Fig. 2P”‘O.
experiments) than when just the -3O-40% of taste buds
on the end of the tongue are engaged30 (as in dorsal-flow
experiments). Furthermore, even when comparing re-
sults from several studies that used the same dorsal-flow
design, the rate of the flow and the temperature of the
solutions can both markedly affect the magnitude of the
result@. It has even been observed that the type of
experimental design can change a mixture interaction
from weakly suppressing to strongly suppressing6, pre-
sumably owing to changes in the shape of the concen-
tration-intensity power functions of the unmixed com-
pound&6,31 (see Fig. 2). In addition, both the experimental
context in which ratings are made and the number of
scales on which a subject rates simultaneously can have
an impact on the magnitude of the ratings”,32-36.
(5) There are very large differences in the types of mixture
interactions depending on whether near-threshold (concen-
trations that are barely detectable or not detectable at all)
or obviously supratbreshold concentrations are examined.
This point is distinct from point 3, which discusses only
391
 x
.ti
2
3
E

,‘-

Concentration

1 Nochang; inslope 1

Enhancement Suppression

,
.
.
_ _ - - .

:.......
......
,i
L

presence
Linear scaling.
concentration-intensity
like a Weibul
of a background
shift the function linearly
The upper

or logistic
Concentration

function
function.
panel depicts

concentration
for a tastant
The lower
a typical

of an added
left or right along the x-axis,
that is shaped
panel shows how the
compound
indicating
Fig. 1
psychometric
roughly

may
that the
1J I
Expansive,
Concentration

linear and compressive portions of a psychometric

Fig. 2

background additive is enhancing or suppressing taste, respectively.

function. All three panels depict the exact same psychometric
Note that the shape of the function, however, is unaltered.
concentration-intensity function for a tastant that is shaped
roughly like a Weibul or logistic function. In the upper panel, the
interactions with easily detected suprathreshold concen-
low-concentration portion of the function is outlined to indicate the
trations of tastants. Subthreshold mixtures of qualitatively
region where the function is expansive. Thus, two solutions of low
different-tasting compounds have generally yielded additive
combinations7, whereas almost all very strongly supra- concentration should combine hyperadditively. In the middle panel,
threshold mixed-quality interactions yield suppression1-4. the moderate-concentration portion of the function is outlined to show
that here it is linear. In this concentration range, two solutions should
In what follows, I review information about various types combine additively. In the lower panel, the high-concentration portion
of mixture interactions between pairs of salty, bitter and of the function is outlined to indicate where it is compressive. Thus,
sour compounds, with an emphasis on the enhancement or two solutions of high concentration should combine hypoadditively.
suppression of taste qualities in binary mixtures. Most
Note that all three of these types of interactions occur whenever the
of the reviewed research focused on single judgements of
function is ‘S’ shaped, and therefore will occur for a function that is
quality magnitude, rather than multiple magnitude ratings
over time, such as in time-intensity experiments. The shifted along the x-axis but maintains its ‘S’ shape as in Figs 1 and 3.
present focus on individual ratings of sourness, bitterness
or saltiness in binary mixtures is not meant to imply that both simultaneous (split-tongue)29 and successive con-
combinations of qualities in mixtures may not elicit new trasts among taste combinations of different qualities.
taste qualities that individual compounds cannot elicit37.
However, many subjects appear to be able to relate to the Mixtures of salts (NaCI) and acids
experimenter’s request to introspect about single attributes Much of the work on salt-acid mixtures is reviewed
of a complex taste stimulus, as they might do during every- in Table 1. The following points briefly summarize the
day experiences, such as when they report that vegetables conclusions that can be drawn from this table.
need salt, or fruit taste too bitter, or lemonade is too sour.
In this review, I will not fully evaluate the literature Subthreshold compounds mixed with subthreshold
regarding the location of mixture interactions (i.e. distin- compounds
guish among effects taking place in the oral epithelium, Salts and acids tended to display slight hyperaddi-
peripheral nerves and the brain). Nor will I review the tivity when subthreshold concentrations were mixed. The
various types of adaptation studies that have examined approximate additivity of these different compounds (for

392 Trends in Food Science & Technology December 1996 [Vol. 71

 and saltiness, respectively, in a variety of media with a
range of methods.

Strongly suprathreshold compounds mixed with strongly

suprathreshold compounds
1 Suppression was much more commonly observed
when strongly suprathreshold concentrations of acids
.--
Change in slope
and salts were mixed. There were also several obser-
,/-.-’ vations of no interaction in mixtures of compounds at
these high intensities. Enhancement was rarely seen.

*‘I
J ,
.,’
__-’ Mixtures of bitter-tasting compounds and acids
Much of the work on bitter-acid mixtures is reviewed
in Table 2. The following points briefly summarize the
conclusions that can be drawn from this table.
Synergy
Subthreshold compounds mixed with subthreshold
compounds
Bitter-tasting compounds and acids tended to display
/, slight hypoadditivity when subthreshold concentrations
were mixed.

Subthreshold (or threshold) compounds mixed with

Concentration suprathreshold compounds
, In one study, Pangbom42 found that caffeine and citric
Fig. 3 acid suppressed each other at very low concentrations.
Nonlinear scaling. The upper panel depicts a typical psychometric In contrast, von Skramlik41 found that the threshold for
concentration-intensity function for a tastant that is shaped roughly
tartaric acid was lowered by quinine HCl. In other
words, sensitivity was enhanced.
like a Weibul or logistic function. The middle panel shows how the
addition of a background concentration of a compound may alter the Weakly suprathreshold compounds mixed with weakly
shape of the function nonlinearly, rendering it steeper or shallower. (or strongly) suprathreshold compounds
The lower panel shows how the presence of the background additive The sourness of citric acid was enhanced by caffeine2,
may shift the function along the x-axis, leftward and steepening it, or whereas the bitterness of quinine sulfate was suppressed
rightward and shallowing it, indicating that the additive is interacting by low concentrations of tartaric acid16.
synergistically or is masking the taste, respectively.
Strongly suprathreshold compounds mixed with strongly
all three mixture types, see also Tables 2 and 3) suggestssuprathreshold compounds
that, at subthreshold concentrations, their effects summate The results with strongly suprathreshold concen-
in a pathway common to both chemicals somewhere in trations of acids and bitter-tasting compounds were
the nervous system, although most likely not in the same mixed. There is an account of sourness being enhanced
peripheral taste receptor cell. by mixture with a bitter compound [H. Gerigk (1955)
iiber die Miigh’chkeiten der Sinnlichen Unterdriickung
Subthreshold (or threshold) compounds mixed with auf dem Gebiete des Geschmacksinnes (On the Possi-
suprathreshold compounds bilities of Sensory Suppression in the Area of the Taste
When subthreshold concentrations of a salt (NaCl) Sense) (PhD thesis), Berlin, Humboldt Universitat,
were mixed with strongly to weakly suprathreshold con- Berlin, Germany], and an account of bitterness being
centrations of various acids, the salt tended to suppress enhanced by mixture with an acid2. On the other hand,
the perceived sourness (or elevated the threshold for the there are two accounts of acids and bitter-tasting com-
acid) (see Table 1). pounds suppressing each other, with the exception that
Similarly, acids suppressed the perceived saltiness of moderate concentrations of tartaric acid are not sup-
salts, with two notable exceptions: subthreshold concen- pressed by quinine sulfate 16,45.Frank et aE.32found that
trations of many acids enhanced the saltiness of NaC13*, citric acid either enhanced or suppressed the bitterness
and tartar% acid lowered the threshold (i.e. enhanced the of quinine depending on how the subjects rated the
saltiness) for NaC141. bitterness.

Weakly suprathreshold compounds mixed with weakly Mixtures of salts and bitter-tasting compounds
(or strongly) suprathreshold compounds Much of the work on salt-bitter mixtures is reviewed
In general, weakly suprathreshold concentrations of in Table 3. To briefly summarize, the following conclu-
acids and salts tended to enhance one another’s sourness sions can be drawn from this table.

Trends in Food Science & Technology December 1996 [Vol. 71 393

 Table 1. Mixtures of salts and acids

Components
Mixture intensity level Salt Acid Outcome Ref.

Subthreshold t subthreshold NaCl Citric Slightly hyperadditive Stevens (1995)’

Subthreshold [or threshold) NaCl Many Subthreshold concentrations of NaCl moderately suppressed Fabian & Blum (1 943)38
t suprathreshold the sourness of acetic, HCI and citric acids without
altering the pH (suppression)

NaCl Many Subthreshold concentrations of NaCl greatly suppressed Fabian & Blum (1943)‘”
the sourness of lactic, malic and tartaric acids without
altering the pH (suppression)

NaCl Many Subthreshold concentrations of many acids increased Fabian & Blum (1 943)38
the saltiness of NaCl (enhancement)

NaCl HCI Seven out of ten panelists reported no interaction; three Fabian & Blum (1 943)38
out of ten reported that subthreshold HCI reduced the
saltiness of NaCl (suppression)

NaCl HCI HCI increased the threshold for NaCl (subtractive mixing) Anderson (1 950)3g

NaCl HCl Thresholds were raised when suprathreshold concentrations Heymans (1 899)40
of one were added to the other (subtractive mixing)

NaCl Tartaric Most subtractive mixtures elevated both thresholds, but tartaric von Skramlik (1962)“’
acid lowered the threshold for NaCl (subtractive mixing)

NaCl Citric Subthreshold concentrations of NaCl reduced the sourness Pangborn (1960)“’
of citric acid (suppression)

Weakly suprathreshold t weakly NaCl Citric Weak NaCl and weak citric acid resulted in enhanced Pangborn & Trabue
(or strongly) suprathreshold saltiness and mixed effects on sourness (enhancement) (1 964)23

NaCl Citric Citric acid enhanced the saltiness of NaCI, when NaCl Kamen et a/. (1961)j
was weak (enhancement); the sourness of weak citric acid
was enhanced by NaCl (enhancement)

NaCl Tartaric High concentrations of tartaric acid enhanced the saltiness lndow (1 969)16
of weak NaCl (enhancement); high NaCl concentrations
enhanced the sourness of weak tartar? acid (enhancement)

NaCl Lactic NaCl enhanced the sourness of weak lactic acid (enhancement) Hellemann (1 992)43

NaCl Acetic NaCl enhanced the sourness of acetic acid in rye bread Hellemann (1 992)43
(enhancement)

,Strongly suprathreshold NaCl Tattaric NaCl and tartaric acid suppressed each other Hambloch & Puschel
t strongly suprathreshold (suppression) (1928)+’

NaCl Tartaric Tartaric acid had no impact on the saltiness of NaCl (no effect) lndow (1 969)16

NaCl Tartaric Tartaric acid and NaCl suppressed each other (suppression) Gerigk (1955)”

NaCl Citric Citric acid reduced the perceived saltiness of NaCl (suppression) Pangborn (1 960)42

NaCl Citric The sourness of citric acid was suppressed by NaCl Pangborn & Chrisp
(suppression); the saltiness of NaCl was enhanced by (1 964)22
citric acid - but variability was very high (enhancement?)

NaCl Citric The saltiness of NaCl was unaffected by citric acid (no effect); Kamen et a/. (1961)*
the sourness of citric acid was suppressed by NaCl (suppression)

NaCl Lactic NaCl suppressed the sourness of lactic acid (suppression) Hellemann (1 992)43

NaCl Acetic NaCl suppressed the sourness of acetic acid (suppression) Hellemann (1 992)43

NaCl HCI NaCl and HCI were mildly enhanced or mildly suppressed Kiesow (1 894)45
depending on the subject (mixed effects)

NaCl HCl The sourness of HCI was unaffected by NaCl (no effect) von Skramlik (1962)41

NaCl HCI NaCl and HCI did not interact with one another (no effect) Bartoshuk (1975)’

NaCl HCI NaCl had no effect on the sourness of HCI (no effect) Cragg (1 937)46

“PhD thesis; see text for details

394 Trends in Food Science & Technology December 1996 [Vol. 71

 [able 2. Mixtures of bitter-tasting compounds and acids

Components

Mixture intensity level Bitter Acid Outcome Ref.

Subthreshold + subthreshold Quinine HCI Citric Slightly hypoadditive Stevens (1995)’

Subthreshold (or threshold) Quinine HCI Tartaric Most subtractive mixtures elevated thresholds, von Skramlik (1 962)4’
t suprathreshold but quinine HCI lowered the threshold for tartaric
acid (subtractive mixing)

Caffeine Citric Subthreshold concentrations of caffeine Pangborn (1 960)42

suppressed the sourness of citric acid
(suppression); subthreshold citric acid suppressed
the bitterness of caffeine (suppression)

tieakly suprathreshold Caffeine Citric Caffeine enhanced the sourness of citric acid, Kamen et a/. (1961)2
t weakly (or strongly) especially at low acid concentrations
suprathreshold (enhancement)

Quinine SO, Tartaric Low concentrations of tartaric acid suppressed lndow (1 969)16
the bitterness of quinine SO, (suppression)

Strongly suprathreshold Variable Variable Bitterness and sourness did not interact Kiesow (1 894)45
t strongly suprathreshold (no effect)

Quinine HCI Tartaric Quinine HCI and tartaric acid suppressed Hambloch & Puschel (1 928)44
each other (suppression)

Quinine HCI Tartaric The sourness of tartaric acid was enhanced Gerigk (1955)’
by quinine HCI (enhancement)

Quinine SO, Tartaric High concentrations of quinine SO4 suppressed lndow (1969)r6
the sourness of low and high concentrations
but not moderate concentrations of tartaric
acid (suppression)

Quinine HCI Citric Citric acid either enhanced or suppressed the Frank et al. (1 993)32
bitterness of quinine depending on how the
ratings were made

Caffeine Citric Citric acid greatly enhanced the bitterness Kamen et al. (1961)*
of caffeine for all concentrations of caffeine
(enhancement)

‘PhD thesis; see text for details

Subthreshold compounds mixed with subthreshold
compounds
Salts and bitter-tasting compounds tended to display
slight hypoadditivity when subthreshold concentrations
were mixed.
Subthreshold (or threshold) compounds mixed with
suprathreshold compounds; weakly suprathreshold
compounds mixed with weakly (or strongly)
suprathreshold compounds; and strongly suprathreshold
compounds mixed with strongly suprathreshold
compounds
All three intensity levels yielded comparable results
and are therefore considered together here. Although
quinine and caffeine are the two most commonly em-
ployed bitter-tasting stimuli in the psychophysical litera-
ture, there is at least one report in which a larger array
of salts and bitter compounds was used’*. In all but one
instance, sodium salts suppressed bitterness to some de-
gree. The opposite was not always true. Although there
are some accounts of bitter compounds suppressing the
saltiness of NaCl, there are others in which saltiness was
not affected2,‘2.47.Notable exceptions to this trend were
MgS04 and amiloride HCl, which were the only two
bitter compounds to suppress the saltiness of NaCl at
moderate salt concentrations12.
The magnitude of the suppression of bitter taste by
NaCl varied greatly for the different bitter-tasting com-
pounds tested, ranging from -5% suppression of the
most bitter concentration of MgSO, tested to -80% sup-
pression of the most bitter concentration of urea. Non-
sodium chloride salts (with the exception of LiCl) were
not able to suppress the bitterness of any compound to
any great degree.
There was a single example of bitterness enhance-
ment by NaCl when mixed with iso-c.u-acids48.
Conclusions
‘How do bitter and sour tastants interact in mixture?’
‘Does sourness increase saltiness when in mixture?’
‘Will bitterness affect saltiness?’
I argue here that such questions do not really make sense.
Mixture interactions will be dependent on the specific
chemicals in question, the magnitude of their perceived

Trends in Food Science & Technology December 1996 [Vol. 71 395

 Table 3. Mixtures of bitter-tasting compounds and salts
I-
Mixture intensity level Bitter Salt Outcome Ref.

Subthreshold t subthreshold Quinine HCI NaCl Slightly hypoadditive Stevens (I 995)’

Subthreshold (or threshold) Caffeine NaCl Subthreshold and threshold caffeine reduced the saltiness of NaCl (suppression); subthreshold Pangborn (I 960)42
t suprathreshold and threshold concentrations of NaCl suppressed the bitterness of caffeine (suppression)
Quinine HCI NaCl Quinine HCI increased the threshold for NaCl (subtractive mixing) Anderson (I 950)39

Weakly suprathreshold t weakly Quinine HCI NaCl Low concentrations of NaCl suppressed the bitterness of quinine (suppression) Hopkins (I 953)47
(or strongly) suprathreshold

Strongly suprathreshold Quinine SO, NaCl All concentrations of NaCl reduced the bitterness of all but the weakest quinine SO, lndow (1 969)16
t strongly suprathreshold solutions (suppression)
Quinine SO, NaCl Quinine SO, had no effect on the saltiness of NaCl (no effect) Hopkins (1 953)47
Quinine SO, NaCl The bitterness of quinine SO, was suppressed by NaCl (suppression) Kroeze (I 980)*8
Quinine HCI NaCl NaCl and quinine HCI suppressed each other (suppression) Hambloch & Puschel (I 928)”
Quinine HCI NaCl Quinine HCI suppressed the saltiness of NaCl (suppression) Cerigk (I 955)”
Quinine HCI NaCl The bitterness of quinine HCI was suppressed by NaCl (suppression) Bartoshuk (I 975)’
Quinine HCI NaCl NaCl suppressed the bitter taste of quinine HCI at the same tongue location (suppression) Kroeze & Bartoshuk (1985)2q
Quinine HCI NaCl NaCl suppressed the bitterness of quinine HCI (suppression) Schifferstein & Frijters (1992)”
Quinine HCI NaCl NaCl suppressed the bitterness of quinine HCI by 41% (suppression) Breslin & Beauchamp (1995)‘*
Caffeine NaCl Caffeine and NaCl had no effect on each other. Although a couple of significant interactions Kamen et a/. (196I)j
were observed, the authors questioned their own findings owing to large error terms (no effect)
Caffeine NaCl NaCl suppressed the bitterness of caffeine by 55% (suppression) Breslin & Beauchamp (1995)12

MgQ NaCl NaCl suppressed the bitterness of MgSO, by 4% (suppression) Breslin & Beauchamp (1995)‘*
Amiioride HCI NaCl NaCl suppressed the bitterness of amiloride HCI by 69% (suppression) Breslin & Beauchamp (1995)i2
KCI NaCl NaCl suppressed the bitterness of KCI by 78% (suppression) Breslin & Beauchamp (I 995)12
Urea NaCl NaCl suppressed the bitterness of urea by 76% (suppression) Breslin & Beauchamp (I 995)12
Urea Na acetate Na acetate suppressed the bitterness of urea by an amount comparable to that of NaCl (suppression) Breslin & Beauchamp (I 995)”
Quinine HCI Na acetate Na acetate suppressed the bitterness of quinine HCI by an amount comparable to that Breslin & Beauchamp (1 995)12
of NaCl (suppression)

Urea Na gluconate Na gluconate was as effective as NaCl at suppressing the bitterness of urea (suppression) Breslin & Beauchamp (I 995)”

Urea LiCl LiCl suppressed the bitterness of urea by 81% (suppression) Breslin & Beauchamp (1 995)12

WQ NaCl MgSO, suppressed the saltiness of moderate concentrations of NaCl (suppression) Breslin & Beauchamp (I 995)i2
Amiloride HCI NaCl Amiloride HCI suppressed the saltiness of moderate concentrations of NaCl (suppression) Breslin & Beauchamp (I 995)12

Iso-a-acids NaCl NaCl enhanced the bitter taste of isoa-acids (enhancement) Yokomukai et a/. (I 994)48

‘PhD thesis; see text for details

 intensity, the particular concentrations examined, whether
they are subthreshold or suprathreshold, and the particular
method that is used to answer the question32*36.
Despite the need for these specificities in order to an-
swer such questions, trends in binary mixture interac-
tions may be seen from the literature. Foremost among
them is that mixtures of subthreshold, threshold and
even weakly suprathreshold compounds behave quite
differently from mixtures of medium to highly supra-
threshold compounds. Mixtures of compounds that elicit
different taste qualities are reported to be additive on
average - usually either slightly hyperadditive or hypo-
additive - when a subthreshold concentration of one is
mixed with a subthreshold concentration of another’.
Subthreshold, threshold and weakly suprathreshold
compounds mixed with threshold and weakly supra-
threshold compounds may result in suppression, but
often instead in enhancement, depending on the par-
ticular compounds in question and their precise inten-
sities (for example, see Ref. 40; also see tables).
Highly suprathreshold mixtures usually suppress each
other, sometimes symmetrically and sometimes very
asymmetrically12~42~49.
The linear decreases (suppression) or increases (en-
hancement) seen in most suprathreshold mixtures are
predicted by the shape of the concentration-intensity
power function for each of the individual compound#.
Compressive power functions for tastants (i.e. those that
have exponents less than one - where doubling the con-
centration results in less than doubling of the intensity)
result in mixture suppression for that compound (it is
less than purely additive). Expansive power func-
tions (i.e. those that have exponents greater than one -
where doubling the concentration results in more than
doubling of the intensity) yield enhancement in mixture
(it is greater than purely additive)31. Note that the effects
of these mixtures are a direct linear prediction from the
shape of the function.
This idea may be extended. Because many psycho-
physical concentration-intensity functions display a sig-
moidal or Weibul-like character, they will, by their very
nature, display both expansion and compression for
lower and higher concentration ranges, respectively (see
Fig. 2). This extension of Bartoshuk’s idea may account
for why salts and acids usually display enhancement at
weak concentrations, when functions are expansive, and
suppression at high concentrations, when functions are
compressive (see Table 1). This idea may also account
for why intense sweeteners often enhance one an-
other-18,19.That is, their mixture interactions are rarely
tested at concentration ranges where the concentration-
intensity functions show saturation (compression), but
rather are tested where the function is expansive (begin-
ning of function) or linear (middle of function), where
enhancement is predicted. Were they to be administered
at the highest testable concentrations, they would be ex-
pected to show suppression. In fact, Frank et aLlo found
that intense sweeteners exhibited hyperadditivity at low
concentrations, additivity at moderate concentrations,
and hypoadditivity at high concentrations.
Trends in Food Science & Technology December 1996 [Vol. 71
Kroeze’” points out that enhancement and suppression
may also often be predicted by the side tastes that many
compounds elicit, such as the sweet taste of weak NaCl
concentrations, which may play a significant role in
suppression or enhancement. For example, the sweet
taste of weak NaCl concentrations may be responsible
for the enhancement of sweetness in mixtures with
sucrose21. Kroezer4 boldly goes on to conclude that be-
cause of the existence of such side tastes, enhancement
in moderate- to high-concentration mixtures is almost
always spurious (with the possible exceptions of
MSG and S-ribonucleotides, certain intense sweetener
mixtures, and NaCl mixed with iso-o-acids). In other
words, the same taste quality is in effect being added
to itself, resulting in apparent ‘enhancement’. Dis-
agreement among laboratories may be accounted for by
differences in the concentrations employed or in the
compounds selected, as just described. However, there
are even simpler arguments to account for disparities in
observations. Pangbom42 suggested that many of the
discrepancies among laboratories may be due to the em-
ployment of expert panels in some studies, and naive
panels in others. Gregson and McCowen50 also attribute
inter-laboratory differences to variations in the sensitiv-
ities of the different populations of panelists. However,
there are several reliable interactions on which different
laboratories concur. The rather impressive interactions
among:
S-ribonucleotides and MSG, which enhance each
other;
certain intense sweeteners, which also enhance each
other; or
sodium salts and certain bitter-tasting compounds, in
which the bitterness is suppressed,
may be separated from all other forms of mixture inter-
action in that these three are reliable, large-magnitude
effects that occur at relatively high concentrations and,
in the case of MSG and ribonucleotides, and certain
intense sweetener mixtures, are not predicted by the
shapes of their power functions6,10J9.Interestingly, it has
been suggested that all three of these interactions have
a peripheral transductive/receptor locus and are not cog-
nitive interactions.
One may surmise functional and adaptive reasons to
explain why the inter-quality interactions discussed here
vary as a function of the different intensity levels. First,
the additivity of compounds at the weakest intensity level
(subthreshold mixtures) has many practical implications.
At the weakest mixture concentrations, when we experi-
ence complex mixtures of subthreshold compounds in our
dally lives, we may use subthreshold additivity effects to
detect their presence. For example, it is possible that many
different mineral waters are distinguishable, despite the
fact that their mineral and salt contents are in the parts
per billion range, because the subthreshold minerals and
salts combine additively to generate distinguishable tastes
(assuming there are no olfactory cues)‘. Second, the di-
versity in mixture interactions at low concentrations,
397
 which generate suppression in some instances and en-
hancement in others, may be utilized to modulate fla-
vors. A good example of this is the role that salt plays in
breads. An acceptable bread does not usually appear to
taste salty, although it contains moderate concentrations
of salt; however, when the same bread recipe is fol-
lowed without the added salt, the bread is markedly
worse in taste, almost unpalatable to most. Glutamate,
and possibly other amino acids, may play a comparable
modulatory role in soup flavor. Third, the high-concen-
tration interactions, which are usually suppressive, may
serve a more general inhibitory function. For example,
many fruit and plants (e.g. citrus fruit, tomato, pine-
apple) are extremely acidic when unripe. As they ripen,
the sugar concentration increases (and the acid concen-
tration may decrease). The increase in sugar concen-
tration serves to suppress the perceived sourness, in the
same way that sugar suppresses the sourness of lemon-
ade. This strategy may allow fruit to inhibit microbial
growth with high acid concentrations, yet retain a de-
gree of attractiveness to certain seed-carrying animals.
Interactions among the three qualities salt, bitter and
sour should be of great interest to the food industry be-
cause they shed light on such major issues as:
l how to modulate or reduce bitterness in foods, oral-
care products and pharmaceuticals;
l how to maintain the effects of NaCl on the flavor of
foods without adding sodium (or while reducing
concentrations);
l how to alter the perception of sourness without chang-
ing the levels of acid present.
Summary
Salts and acids
Interactions among salts and acids were non-mono-
tonic over different concentration ranges. Subthreshold
salt concentrations tended to suppress sourness and vice
versa. However, there are two examples of subthreshold
concentration of acids enhancing saltiness. Moderately
weakly suprathreshold concentrations of acids and salt
tended to enhance one another. Strongly suprathreshold
concentrations of salts and acids tended to suppress one
another.
Bitter-tasting compounds and acids
Interactions among bitter-tasting compounds and
acids were also non-monotonic over different concen-
tration ranges, and seemed particularly variable across
laboratories. There is some indication that weakly supra-
threshold sourness is enhanced by bitterness but that
weakly suprathreshold bitterness is suppressed by sour-
ness. At higher concentrations, acids and bitter compounds
either suppressed or enhanced one another depending on
the stimuli in question and the experimental methods
employed.
Salts and bitter-tasting compounds
Sodium salts almost universally suppressed per-
ceived bitterness across a wide range of bitter-tasting
398
compounds and concentrations. However, the magnitude
of the bitterness suppression varied greatly from com-
pound to compound, and one bitter compound was en-
hanced by NaCl. Non-NaCl salts did not tend to have as
great a general suppressive capacity, with the exception
of LiCl. Only two bitter-tasting compounds suppressed
the saltiness of NaCl clearly.
Acknowledgements
I wish to thank Gary K. Beauchamp and Bruce P.
Halpem for reading and commenting on an earlier draft
of this manuscript.
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effectively used to mask bitter taste are discussed.
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Evidence linking sweet and bitter tastes
Structure-taste relationships
The relationship between chemical structure and taste
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very closely related. Small changes in structure can con-
D. Eric Walters is at the Department of Biological Chemistry,
University of Health Sciences/The Chicago Medical School, 3333
Bay Road, North Chicago, IL 60064, USA (fax: tl-847-578-3240; e-mail:
walterse@mis.finchcms.edu).
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Special Issue on Flavour Perception
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How are bitter and
by
sweet tastes related?
D. Eric Walters
vert sweet compounds to bitter ones, and vice versa, as
shown in Fig. 1. ‘Sugar’ and ‘sweet’ are practically syn-
onymous, yet many sugar esters and halogenated sugars
are intensely bitter. Other sugar derivatives, such as the
chloro-sugar sucralose, have sweetness potencies that
are hundreds of times that of sucrose. Numerous classes
of structures (e.g. peptides, oximes, terpenes, ureas,
guanidines and saccharin derivatives) include examples
that taste sweet, examples that taste bitter, and examples
that taste both bitter and sweet’. Neohesperidin is a bitter
terpene glycoside that is found in citrus peel; cleavage
of a single carbon-oxygen bond produces the intensely
sweet compound neohesperidin dihydrochalcone.
Finch Sensory experiments
Green Sensory experiments also point towards a connection
between bitter and sweet tastes. Mixture suppression is
commonly observed: when sweet and bitter compounds
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