Physiology & Behavior, Vol. 21, pp. 609-613. Pergamon Press and Brain Research Publ., 1978.

Printed in the U.S.A.

Sweet Taste of Dilute NaCl: Psychophysical

Evidence for A Sweet Stimulus I
L I N D A M. B A R T O S H U K , C L A I R E M U R P H Y A N D C A R O L T. C L E V E L A N D

John B. Pierce Foundation Laboratory and Yale University, N e w Haven, CT 06519

( R e c e i v e d 15 M a y 1978)

BARTOSHUK, L. M., C. MURPHY AND C. T. CLEVELAND. Sweet taste of dilute NaCl: Psychophysical evidence for
a sweet stimulus. PHYSIOL. BEHAV. 21(4) 609--613, 1978.--Dilute NaCI tastes sweet. This sweetness could result from
ceding confusions in the nervous system such that weak NaCI produces neural signals resembling those for sweeteners like
sucrose. On the other hand, an analysis of the structural chemistry of water-salt interactions suggests that water shells
organized around cations may actually provide a sweet stimulus indistinguishable (to the receptor molecules) from more
conventional sweet stimuli. In the present experiment, the sweetness of weak NaCl was abolished in two ways: by
adaptation to sucrose and by topical application ofGymnema sylvestre. Since these two operations also abolish the sweet
taste of a variety of conventional sweetners, the sweet taste of NaCI appears to result from the presence of a sweet
stimulus.

Taste Sweet NaC1 Gymnemasylvestre Adaptation Water

SKRAMLIK [49] demonstrated that the tastes of a variety of clusions. Several lines of evidence suggest that saltiness
salts could be matched with mixtures of NaCI, glucose, tar- should be attributed to cations rather than anions. Elec-
taric acid, and quinine. Since salts dissociate in water to a trophysiological analyses of taste responses in rats led Bei-
mixture of cations, anions, and undissociated molecules, dler [10-12] to conclude that the cations of simple inorganic
several early psychophysical studies were oriented toward salts excite taste impulses when they bind to receptor sites
determining the contribution of each component to the total on the taste membrane while the anions inhibit the mag-
taste of the salt [18, 24, 25, 28, 45]. Saltiness was usually nitude of response produced by the cations. Other transduc-
attributed to the anions, particularly the chloride and sulfate tion models for salts also identify the cation as the part of the
anions since among salts with a common cation, the chloride salt responsible for stimulating nerve impulses [15,26].
and sulfate salts were usually the saltiest. Bitterness was Cross-adaptation studies, done both electrophysiologically
attributed to both cations and anions. For example, the bit- [51] and psychophysically [36,52] tend to show that salts
terness of KCI was attributed to the potassium cation and the with common cations produce the best cross-adaptation.
bitter taste of NaI was attributed to the iodide anion because Stimulation of the rat tongue with anodal current (which
NaCI is not bitter. Bitterness was also attributed to a variety moves cations toward the tongue) produced neural re-
of large anions because among salts with a common cation, sponses similar to those produced by salts [50]. These lines
salts tend to get increasingly bitter as the anion gets larger of evidence are consistent with the attribution of saltiness to
[40]. Sourness was attributed to acid salts [40]. Sweetness the cation. The psychophysical data on saltiness can be rein-
was attributed to both cations and anions. For example, be- terpreted in the light of this evidence and the suggestions of
ryllium salts including BeCh are sweet. By the same logic Beidler [10] and DeSimone and Price [15] on the role of
that attributed the bitterness of KCI to the potassium cation, anions. Salts with chloride and sulfate anions will be the
the sweetness of BeCI~ was attributed to the beryllium ca- saltiest because these anions interfere less than larger anions
tion. The sodium salt of saccharin is an example of a sweet with the taste of the cations. The logic used in the
salt in which the sweetness was attributed to the anion. psychophysical studies was based on the assumption that the
Renqvist [45] called attention to a sweet taste associated contributions of the different components of a salt are addi-
with some dilute salts including NaCI that could not be at- tive. If instead, anions interfere with cation taste, then the
tributed to cations, anions, or undissociated molecules. The conclusions of the psychophysical studies cannot be correct.
sweet taste of dilute salts has subsequently been observed in New insights into the nature of water-salt interactions as
a variety of studies [1, 8, 13, 19, 28]. Renqvist [45] attributed well as the structural chemistry of sweet compounds has led
this sweet taste to hydroxyl ions that he believed were pro- to the rejection of Renqvist's suggestion that the sweet taste
duced by a hydrolysis reaction between water and salt. of dilute salt is produced by hydroxyl groups formed by the
There are now reasons to revise many of the above con- hydrolysis of water [18] and replaced it with the suggestion

aWe thank H. L. Meiselman for the purified Gymnema sylvestre extracts.

C o p y r i g h t © 1978 Brain R e s e a r c h P u b l i c a t i o n s Inc.--0031-9384/78/100609-05502.00/0

 610 BARTOSHUK, MURPHY AND CLEVELAND

that the sweet taste is produced by the water of hydration of METHOD

cations [16, 18, 46]. According to tlalS view, water shells lbrm
around cations such that the structure of the water shell Subjects
conforms to the AH, B structure suggested by Shallenberger Five males and five females, all of whom had previously
and Acree [46] as that common to all sweet substances. served in psychophysical experiments, served as subjects.
Two theories of the neural coding of taste equality suggest The data from an additional 3 subjects were deleted from the
additional considerations about the attribution of qualities to data analysis because these subjects either did not taste the
components of salt solutions. The across-fiber patterning sweet taste or the salty taste of NaCI in the concentration
theory [20-22, 41, 42] and the labeled-line theory [43,44] can range tested.
explain the sweet taste of dilute NaC1 as a coding confusion.
According to the across-fiber patterning theory, two stimuli Stimuli
producing similar patterns of activity across fibers will have
similar tastes. The sodium cations of dilute NaC1 might bind Aqueous solutions of reagent grade sucrose (0.032 M) and
to their customary receptor sites in the normal way but gen- NaCI were prepared with dionized water. Nine concentra-
erate a profile more similar to that produced by sucrose tions of NaCI ranged from 0.000032 M to 0.32 M and were
than that produced by NaCI. In such a case, dilute NaC1 evenly spaced half-log steps apart on a logarithmic scale
would taste like sucrose. According to the labeled-line (i.e., this means that they differed from each other by a
theory a fiber that codes sweetness does so no matter what factor o f about 3.2 on a linear scale). Solutions were warmed
the stimulus actually is. The sodium cations of dilute NaC1 to 34 ° (approximately the temperature of the extended
might bind to their customary receptor sites but if those sites tongue). Gymnema sylvestre was purified by the method of
were on cells synapsing with fibers coding sweetness, the Dateo [5,14].
taste quality evoked would be sweet. Note that none of the
proposed explanations for the sweet taste of dilute NaCI Apparatus
account for the fact that the effect occurs only for dilute Stimuli were delivered through a McBurney flow system
NaCI. This will be discussed further below. [33] at a constant rate of 4 ml/sec.
The present study uses cross-adaptation and the effects of
Gymnema sylvestre to distinguish between the coding con- Procedure
fusions described above and the presence of a sweet stimulus
as causes for the sweet taste of NaCI. Exposure of the Preliminary testing was carried out with each subject to
tongue to sucrose diminishes or abolishes the sweet taste of determine adaptation time for 0.032 M sucrose flowed over
sucrose (self-adaptation) and also the sweet tastes of a vari- the tongue. Seven of the 10 subjects adapted completely
ety of other substances (cross-adaptation). This suggests that within 20 sec. The remaining 3 subjects took from 50 see to 2
these substances produce their sweet tastes via the same min to adapt. Since rinses of this length were not feasible,
receptor mechanism [32]. Topical application of Gymnema the rinse time for all conditions was held constant at 20 sec.
~ylvestre abolishes the sweetness of sucrose as well as a The experiment consisted of 2 parts run in a single session
variety of other sweet substances [5, 6, 27, 39, 47, 54] with approximately I hr in length. In part 1 there were two condi-
the exception of chloroform [29]. Gymnema sylvestre is be- tions, one in which a 20 sec water rinse was flowed over the
lieved to affect sweetness by somehow preventing normal subject's tongue before each stimulus, and a second in which
binding of sweet stimuli to their receptor sites [17,23]. The the rinse was 0.032 M sucrose. The 11 stimuli (9 concentra-
coding confusion explanations for the sweet taste of dilute tions of sodium chloride, 0.032 M sucrose, and deionized
NaCI are based on the normal binding of NaCI to its receptor water) were presented once under each condition in a ran-
molecules. Since sucrose adaptation and Gymnema sylvestre domized order. Part 2 consisted of a single condition in
affect the binding of sucrose to its receptor molecules, aboli- which the subject's tongue was treated with Gymnema syl-
tion of the sweet taste of dilute NaC1 by sucrose adaptation vestre: The subject sipped the extract and retained it in the
and treatment with Gymnema sylvestre would be consistent mouth for 60 sec. This treatment was repeated after either
with the presence of a sweet stimulus (i.e., a stimulus that the fifth or the sixth stimulus. Each of the 11 stimuli used in
binds to the same receptor molecules as sucrose) in dilute Part 1 was presented once in randomized order during Part 2
NaC1. and was preceded by a deionized water rinse. The entire
There is some evidence that sucrose can cross-adapt the experiment was repeated for each subject.
sweet taste of weak NaCI. McBurney [32] showed that adap- Throughout the experiment, the subject sat before the
tation to 0.32 M sucrose decreased the sweet taste of 0.01 M delivery tube of the flow system. When instructed to do so,
NaCI and McBurney and Bartoshuk [34] showed that adap- the subject extended his/her tongue into the flowing
tation to 0.18 M sucrose decreased the sweet taste of 0.01 M stimulus. After the presentation of the rinse and the
NaCI (Walsh test, p =0.05). However, there is an alternative stimulus, the subject gave magnitude estimates o f the inten-
explanation for this apparent cross-adaptation. Adaptation sities of the qualities o f the stimulus [52,53].
to sucrose tends to make water taste bitter [3,37] and this
bitter taste might suppress the sweet taste of weak NaC!
Data Analysis
through a mixture interaction [2]. The present study elimi- To correct for differences in moduli used by the subjects,
nates this possibility by using a lower adapting concentration the data were normalized in the following way: for each sub-
of sucrose (0.032 M). ject the geometric mean of the magnitude estimates of the
If Gymnema sylvestre and cross-adaptation with sucrose intensities of 0.032 M, 0.1 M, and 0.32 M sodium chloride
abolish the sweet taste of NaCI then the sweetness of weak following the water rinse was set equal to 100 and the remain-
NaC1 cannot be explained as a coding confusion by either ing responses multiplied by the appropriate factor. Because
coding theory. Rather, the sweet taste of NaCi must then be of the presence of 0 judgments in the data set, arithmetic
produced by the presence of a sweet stimulus. means were used in the remaining stages of data reduction.
SWEET TASTE OF D I L U T E NaCI 611

200 ADAPTATION TO
WATER
ADAPTATION TO
.052 M SU
I TOPICAL APPLICATION
OF
I
GYMNEMA
~,. 15o

I00 -

50-

I.. O: i - _

.20.060, o6, ;,, - ] ,20

" ' '.ooo, .oo,
' '
.o, i .20 .oool .oo, .o, .,
MOLAR CONCENTRATION NoCI

FIG. 1. Magnitude estimates ( - 1 SE) of the perceived intensities of the saltiness, sweetness, sour-
ness, and bitterness of NaCi after adaptation to water, adaptation to sucrose, and topical application of
Gymnema sylvestre.

The arithmetic mean for the 2 responses to each stimulus after water and after sucrose adaptation, p<0.01) but not
was calculated for each stimulus for each subject. The arith- after Gymnema sylvestre (p<0.3). The small increase in the
metic mean was then taken across subjects for each stimulus saltiness of NaCl after adaptation to sucrose and Gymnema
and the standard errors of these means were calculated. sylvestre was not statistically significant (Friedman two-way
analysis of variance on the differences between saltiness
after water and after Gymnema sylvestre, p>0.2).
RESULTS
Figure 1 shows the differences in the magnitude estimates
of the four qualities produced under the three different con- DISCUSSION
ditions. Both adaptation to 0.032 M sucrose and topical The results of this study are consistent with the sugges-
treatment with Gymnema sylvestre suppressed the sweet tion that dilute NaCI actually contains a sweet stimulus that
taste of the low concentration of sodium chloride. These interacts with the same receptor molecules as sucrose. The
effects were tested with the Friedman two-way analyses of possible explanations of the sweet taste of weak NaCI based
variance [48] applied to the sweetness scores for the water on the coding confusions between salty and sweet that were
adaptation condition and applied to the differences between discussed above are not supported by these results. Note
the sweetness scores for the water and sucrose adaptation that the failure of the present data to support these interpre-
conditions and for the water adaptation and Gymnema syl- tations is not evidence against either coding theory.
vestre conditions (p<0.001 for all 3 analyses). It is tempting to view the perceived intensity of the sweet
The effectiveness of sucrose adaptation was checked by taste of weak NaCl as a reflection of the concentration of
including sucrose itself as a stimulus in the experiment. The sweet stimulus present; however, this is unwarranted. The
sucrose concentration used as the adaptation stimulus (0.032 sweetness of NaCl begins to decline at about the concentra-
M) produced a sweetness of 45.1 _+ 6.0 when tasted as a tion at which saltiness begins to climb. The decline in sweet-
stimulus but after 20 sec of adaptation to itself, 0.032 M ness cannot be used as an indication of the actual concentra-
sucrose was judged 5.4 -+ 3.1 sweet. After treatment with tion because the decline may be the result of mixture sup-
Gymnema sylvestre, 0.032 M sucrose was judged 0.02 -+ 0.2 pression, that is, the more intense saltiness might be ex-
sweet. Adaptation to 0.032 M sucrose did not produce a pected to suppress the weaker sweetness [9].
significant bitter water taste (2.06 _+ 1.66 for water after suc- In general, modern psychophysical studies show that
rose compared to 0.64 +_ 0.64 for water after water). substances with common taste qualities cross-adapt one an-
Sucrose adaptation and Gymnema sylvestre produced other [32, 36, 38, 52] while substances with different taste
some small changes in other qualities in addition to the re- qualities do not [34,35]. The major exception to this is the
duction in sweetness. The sourness of NaCI was significantly failure of some bitter stimuli to cross-adapt [34,38]. Contrast
more intense after adaptation to sucrose (Friedman two-way or cross-enhancement phenomena, in which exposure to a
analysis of variance on the differences between sourness substance of one taste quality is supposed to increase the
612 BARTOSHUK, MURPHY AND CLEVELAND

i n t e n s i t y o f a n o t h e r t a s t e quality, h a v e b e e n a t t r i b u t e d to altered w h e n t h o s e s u b s t a n c e s are mixed. T h e i n t e n s i t i e s c a n

w a t e r t a s t e s [34,35]. T h a t is, e x p o s u r e to t h e first s u b s t a n c e b e i n c r e a s e d or d e c r e a s e d d e p e n d i n g at least in p a r t o n t h e
m a k e s w a t e r t a k e o n a t a s t e [1, 3, 7, 8, 31, 33, 37] so t h a t t h e shapes of the psychophysical functions of perceived inten-
t a s t e o f t h e s e c o n d s u b s t a n c e b e c o m e s t h e s u m o f the solute sity v e r s u s c o n c e n t r a t i o n [2,4]. A d a p t a t i o n to o n e c o m p o -
t a s t e a n d the w a t e r s o l v e n t taste. n e n t of a m i x t u r e c a n e l i m i n a t e m o s t o f t h e m i x t u r e i n t e r a c -
I n the p r e s e n t s t u d y , t h e r e are s o m e a p p a r e n t v i o l a t i o n s t i o n s in t w o - c o m p o n e n t m i x t u r e s ( B a r t o s h u k , u n p u b l i s h e d
o f t h e a b o v e g e n e r a l i z a t i o n s t h a t d e s e r v e c o m m e n t . T h e in- d a t a ; [30]). In t h e p r e s e n t e x p e r i m e n t , a n i n c r e a s e in t h e
c r e a s e s in t h e saltiness a n d s o u r n e s s o f N a C I as a result o f s o u r n e s s a n d t h e saltiness o f N a C i following s u c r o s e a d a p t a -
a d a p t a t i o n to s u c r o s e c a n n o t b e a t t r i b u t e d to w a t e r t a s t e s tion c o u l d b e t h e result o f a r e l e a s e o f m i x t u r e s u p p r e s s i o n
a n d so m i g h t s e e m to r e p r e s e n t g e n u i n e c a s e s o f c r o s s - p r o d u c e d b y s u c r o s e . T h e fact t h a t t r e a t m e n t with Gymnema
e n h a n c e m e n t ; h o w e v e r , t h e r e is a n o t h e r m a j o r t a s t e p h e - sylvestre a n d a d a p t a t i o n to s u c r o s e p r o d u c e d similar small
n o m e n o n that m u s t b e c o n s i d e r e d : t a s t e m i x t u r e i n t e r a c - i n c r e a s e in the saltiness a n d s o u r n e s s o f NaC1 s u p p o r t s this
tions. T h e t a s t e i n t e n s i t i e s p r o d u c e d b y s u b s t a n c e s c a n b e idea.

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