1-176
Session VI. Factors in Reducing Salt Intake
High Salt Intake
Sensory and Behavioral Factors
Gary K. Beauchamp and Karl Engelman
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Salt (NaCI) is a ubiquitous component of diets in developed countries. A major reason for this
is that people judge many salted foods as more palatable than the same foods without salt
Because recent evidence indicates that an acceptable salt substitute is unlikely, an understanding of the behavioral and sensory factors involved in maintaining high salt preference is a
prerequisite to successful programs aimed at reducing intake. Although little evidence exists for
a genetic determination of individual differences in consumption and preferred level of salt,
more research in this area is necessary. Considerable data support the view that the optimal
level of salt in the diet is determined in part by the level an individual is currently consuming;
increasing or decreasing customary salt intake, as long as the salt is tasted, increases or
decreases the preferred level of salt in food. Although these data are consistent with a
hypothesis that optimal salt preferences are learned, other data, from both animal models and
human developmental studies, suggest that salt preference has an innate component Furthermore, early experience with low or high salt diets may have a long-term impact on preferred salt
levels. Liking for salt, similar to liking for sweets, has an innate basis that can be modified by
individual experience. (Hypertension 1991;17[suppl I]:I-176-I-181)
S
ubstantial evidence from epidemiological and
experimental sources links dietary NaCI (salt)
intake to hypertension, as has been reviewed
elsewhere.1 For this reason, reduced-salt diets are
frequently recommended for individuals with hypertension2 and have also been recommended for the
US population as a whole.3 Here, we discuss selected
sensory factors that must be considered in developing
successful programs to limit or decrease intake of
salt.
Because salt is required for life, it is not surprising
that regulatory systems have evolved to ensure discovery, recognition, and consumption of sufficient
sodium to meet that requirement. A focal point in
these systems is the sense of taste. Salty taste is one
of a relatively small number of primary taste qualities, others being sweet, bitter, sour, and perhaps a
few others.4
Humans express preferences for salty foods over
those same foods without salt. Factors responsible
for the preference are discussed below and are
complex. However, as a consequence of this preferFrom the Monell Chemical Senses Center (G.K.B.) and the
Hypertension Section and Clinical Research Center (K.E.), Hospital of the University of Pennsylvania, Philadelphia, Pa.
Supported by grants HL-31736 and DC-00882 from the National
Institutes of Health and by Clinical Research Center grant RR00040.
Address for correspondence: Gary K. Beauchamp, Monell
Chemical Senses Center, 3500 Market Street, Philadelphia, PA
19104-3308.
ence, many of the foods available for purchase already contain relatively high levels of salt. Recent
research indicates that perhaps less than 5-10% of
the average sodium consumption comes from what is
generally termed discretionary sources (salt shakers
and salt added during home cooking).5 Consequently, following advice to lower sodium intake by
ceasing to use discretionary salt is likely to result in a
very small overall decrease in sodium intake. For
there to be a substantial decline in sodium intake on
a populationwide basis, a reduction in the amount of
salt in prepared foods, both at the store and sold in
restaurants, and a general decrease in consumption
of very high salt foods appears to be necessary.
That most adults prefer salty foods rather than
pure salt itself has a methodological implication for
research designed to understand factors responsible
for high salt intake. To evaluate the pleasantness of
the taste of salt, it must be added to a familiar food.
Salt taste preference studies that use saline solutions
are generally very difficult to interpret.
Salt Substitutes?
A series of recent observations leads to the conclusion that it is very unlikely that a practical salt
substitute can be developed in the near future.
Although there are now known to be a large and
growing number of sweet-tasting substances, many of
them rather similar to the taste of sucrose, believed
to be the purest sweet, very few compounds taste
predominantly salty (Table 1). This suggests that a
Beauchamp and Engelman Factors Controlling High Salt Intake
TABLE 1. Comparison of Numbers of Sweet- and
Salty-Tasting Compounds
Sweet compounds (partial list)
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Acesulfame-K
Lo han fruit
Alitame
Maltitol
2^-Anhydro-D-mannitol
Mannitol
Aspartame
3-0-Methyifructosc
Chlorinated sucroses
Miraculin
Chlorogenic acid
Monellin
Cyclamate
Osladin
Dihydrochalcones
Perillartine
Dulcin
Phyllodulcin
Fructose
Rebaudioside A
Glucose
Saccharin
Gfycyrrhizin
Sorbitol
High fructose corn syrup
Stevia
Hydrogenated glucose syrup
Stevioside
Isomalt/palatinit
Trihalogenated benzamides
L-sugars
Tryptophan and derivatives
Lactitol
Xylitol
Salty compounds
NaCl
ua
KCI*
Monosodium glutamate*
*Salty but with other tastes sometimes predominant
random search for non-NaCl salty compounds is
unlikely to be successful.
Recent advances6-9 in the understanding of the
mechanisms of sweet and salty tastes may help to
explain the very small number of substances that taste
salty. Sweet receptors are thought to be protein-based
and likely respond to some aspects of molecular shape.
As long as a molecule has the appropriate conformation, according to this view, it will taste sweet. In
contrast, it is thought that at least one component of
salt taste reception involves passage of the sodium ion
through sodium channels at the apical end of the taste
bud. The major evidence for this comes from electrophysiological6 and biophysical7 studies in experimental
animals demonstrating that the diuretic amiloride reduces salt stimulation by about 50%. Whether an
equally robust effect exists in humans is unclear,8-9 and
more studies are needed to clarify this issue. Since the
sodium channels are very specific, it may be exceedingly
difficult to find a nonsodium, nonlithium substitute to
mimic salty taste. Although a salt enhancer may be
possible, the use of substances that are involved in
opening and closing sodium channels as food additives
is problematic.
If it is unlikely that the sensory attributes of salt
can be mimicked by a nonsodium, nonchloride substitute, as is happening with the use of aspartame as
an artificial sweetener, reduction of salt intake will
require other strategies. The development of other
strategies involves an understanding of the determinants of human sensory responses to salt, including
1-177
knowledge of the forces that influence preferences
for high salt levels in many foods.
Determinants of Human Salt Preference
For heuristic purposes, the following presentation
divides discussion of the reasons for high salt intake
into physiological, genetic, psychological, and developmental categories. We recognize that, in reality,
these categories of explanation overlap and are not
mutually exclusive.
Physiological Factors
Most adults in the United States consume 6-12 g
NaCl/day (100-200 meq), an order of magnitude or
more above published requirements.10 Although it
would thus appear unlikely that this level of consumption actually reflects some sort of physiological
need, that has in fact been proposed,1112 despite the
fact that some people live on as little as 20 meq/day
or less without apparent harm.13 At the extreme, the
Yanomamo Indians are reported to consume approximately 1 meq sodium/day, as determined by urinary
excretion (extra loss in sweat is possible), and other
unacculturated peoples also have been reported to
chronically consume very low sodium diets.14 Indeed,
there is exceedingly little human evidence that even
an extremely low sodium diet and sodium depletion
are followed by a greatly increased desire for salt or
what has been termed "salt appetite," as distinguished from the "salt preference" (the choosing of
salty food even when sodium is not needed) observed
in animals and humans who are sodium replete.
In a recent study,15 we placed human volunteers on
a very low sodium diet (5-10 meq/day) and treated
them with diuretics to further deplete them of sodium. They did not express a strong hunger for salt;
this finding is in basic agreement with several earlier
studies.16 However, when asked to rate how much
they would like to eat each of the 29 foods varying in
salt content, the sodium-depleted subjects did express a greater desire for salty foods, and in taste
tests they tended to prefer higher levels of salt in
food while depleted. These data, albeit suggestive, do
not conclusively demonstrate a physiologically determined salt appetite in adult humans since a psychological explanation is also possible: extremely low
sodium diets are bland and unpalatable when contrasted with the previously experienced food, and it
may not be the salt taste these subjects desire so
much as the more flavorful food associated with
saltiness. However, since preference for sweet foods
and sucrose in food declined during depletion, a
simple desire for more orosensory stimulation cannot
explain these results. This issue needs further investigation. A finding that increased salt preference
among sodium-depleted subjects is due to psychological factors would further argue against the need
hypothesis.
Related work17 has shown that normal subjects,
when administered a hydrochlorothiazide diuretic,
exhibited a compensatory increase in sodium intake.
1-178
Supplement I Hypertension Vol 17, No 1, January 1991
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Subjects expressed no obvious desire for increased
salt, and in fact, it was not possible to determine the
source of the increased intake. Interestingly, this
effect was not observed with the diuretic amiloride,
so it would seem that simple sodium loss could not
account for the observed results.
Sodium depletion is generally seen as the adequate
and necessary condition for salt appetite or enhanced
salt preference. Depletion of other nutrients may
also stimulate avid salt consumption. Tordoff et al18
found that calcium-deficient rats increased voluntary
sodium intake. It is possible that mineral deficiencies
in general may stimulate salt intake because there
are no specific tastes for other minerals and because,
in nature, salty tasting substances are almost always
associated with other minerals. Perhaps in an analogous fashion, protein deficiency could stimulate salt
appetite since protein-rich animal foods contain relatively high amounts of sodium. Torii et al19 and
Kimura20 have found that very low protein diets or
diets unbalanced in amino acids stimulate dramatic
increases in salt consumption in rats. This is attributed to the effects of altered protein diets on fluid
balance.
It is generally believed that individuals in cultures
in which a primarily vegetarian diet is consumed
need and crave more salt than do primarily meateating peoples.21 The traditional explanation has
been that meat-eaters obtain substantial amounts of
sodium in meat.22 Although it is certainly true that
meat diets generally contain more sodium than vegetable diets, there is little evidence that the low
sodium diets that might exist in vegetarian cultures
would lead to an elevated desire for salt. Vegetarian
diets in native populations are not only low in salt but
may also be low in protein; perhaps protein content,
rather than sodium content, of the diet underlies an
increased desire for salt, or perhaps both protein and
sodium content influence this desire.
Genetic Factors
Genetic factors have been proposed to account
for differences between groups and individuals in
their salt requirements, intakes, and preferences,
but the evidence favoring that explanation is not yet
strong. For example, it has been speculated that
blacks may have a different mechanism than do
whites for handling sodium. In particular, they may
be less efficient in dealing with excess salt.23 This
could contribute to the high prevalence of hypertension among blacks when salt is readily available.24 There is a single report23 that black adolescents and adults prefer higher levels of salt than do
whites, although recent studies with children did
not confirm this.26 Additionally, it is problematic
whether genetic differences would necessarily underlie a race difference in preference.
Two twin studies have failed to find a heritable
component to salt taste preference. These studies
cannot be considered definitive, however. In one,27
sensory testing was brief and perhaps inappropriate
because it used salt solutions rather than foods to
assess preference. In our twin study,28 the use of very
small numbers of twin pairs rendered the negative
results difficult to interpret. Consequently, a genetic
explanation for individual differences in salt taste
preference remains a possibility, especially in view of
species and strain differences that have been observed in nonhuman animals.22 More work is needed
both in human populations and in animal models.
Psychological Explanations
Experimental studies have demonstrated that if
salt intake is reduced by 30-50% over a period of
time, the optimal level of salt in foods declines. The
amount of salt required to optimize food flavor is
decreased to about 65-75% of the prediet amount.29
Importantly, this change is gradual, taking probably
1-2 months to occur.
For several reasons, it is most likely that this
change is due to the altered (decreased) experience
of tasting salty foods rather than a physiological
response to the change in the amount of sodium
consumed. First, the gradual nature of the effect, as
noted above, would suggest that experience plays the
major role. Second, we found that if salt consumption
was increased by requiring subjects to add approximately 10 g salt to their food, preferred levels of salt
in food (but not in aqueous solution) increased.
However, if the same amount of additional salt was
consumed as tablets, and thus not tasted, there were
no changes in taste preferences.30 Apparently, a
sensory adaptation to different levels of salt accompanies a change in salt intake. To some extent,
people like the level of salt they taste rather than, or
in addition to, choosing the taste level of salt they
like.
The third reason for believing this change in
optimal salt level after dietary change to be a psychological phenomenon comes from another study,31 in
which we reduced salt in the diet of subjects by 50%
but allowed them continued ad libitum use of table
salt. Under these conditions, their use of added salt
increased almost fourfold but did not approach compensation for the amount removed. In fact, they only
replaced 20% of the decrement; thus, there was an
overall decrease of approximately 40% in sodium
intake (Figure 1). In this instance, we suggest that
salt was added "to taste"; this finding further implies
that although the salt content of their regular food
was in a range that the subjects found palatable, it
was unnecessarily high, probably because it was
dispersed throughout the food, rather than all being
on its surface, and easily accessible to the taste
receptors. This study suggests a novel technique to
reduce salt intake, in that individuals were able to use
as much salt as they wished while still reducing total
salt intake. For this to be practical, however, it will be
necessary for there to be wide availability of all
categories of lowered sodium foods. In terms of
control of salt preference, the importance of this
research is that there were no changes in taste
Beauchamp and Engelman
250
200
Factors Controlling High Salt Intake
1-179
associated with alterations in the amount of NaQ
available to the body. Although more extreme depletions may raise salt preferences as described above,
the extent to which this is due to depletion rather
than the blandness of the foods is unknown.
Developmental Factors
4
5
6
7
8 9
Weeks
10 11 12 13 114
18
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FIGURE 1. Graph showing mean daily excretion of sodium
and mean daily amount of sodium ion used from salt shakers
in subjects when they were fed at the Clinical Research Center
(dotted area) and when they were on their usual ambulatory
diets (weeks 1 and 18). All points represent 11 subjects, except
for week 18 (n=8). *Significant difference between comparison weeks. Reprinted from Reference 29 with permission.
preference, even though salt intake was decreased
40%. We suggest that subjects, by adding salt to the
outside of their food, obtained sufficient salty-taste
experience to prevent any preference changes.
In sum (Figure 2), these studies support the conclusion that changes in salt taste preferences after
halving or doubling salt intake are mediated by
hedonic/cognitive expectations based on current dietary experience and not by physiological changes
No
100T
1I
50
u
nCTBOMQ NO
TobM
i
CD
- 1 0 0 *•
FIGURE 2. Bar graphs showing changes in sodium intake as
determined by 24-hour urine values and changes in salt taste
preference (most preferred concentration of salt in soup) in
three experiments. Bars with asterisks indicate significant
(p<0.05) change from baseline. Left panel: Moderate sodium
reduction is followed by a decrease in optimal salt levels
(Reference 27). Middle panel: More extreme sodium restriction with diuretic treatment induces a moderate increase in
optimal salt levels (Reference 29). Right panel: An increase in
sodium intake, but only if tasted, also induces a moderate
increase in optimal salt levels (Reference 28). Not shown are
the differing time courses for these taste changes. Effects seen
on the left panel occur gradually over 1-3 months, whereas the
changes seen on the other two panels are evident within days
or a few weeks after the treatment.
It is in infancy and childhood that the strongest
claims for an experiential influence on salt taste have
been made.32 However, there are conflicting and
confusing data concerning the early development of
salt preference and how experience may serve to
modify or perhaps permanently establish preference.
Newborn infants either are indifferent to33 or
avoid34 moderate to high concentrations of saline
solution relative to water. By the time children are
2-3 or more years of age, preferences for salty foods
over those same foods without salt are common.26
The juxtaposition of these two observations has led
investigators to conclude that salt preference is
learned, although that conclusion does not necessarily follow. We found that a preference for salt
solutions over plain water, although not evident in
infants less than 4 months of age, was evident in
infants 4-23 months of age. This was observed in
three separate cross-sectional studies, all using a
brief presentation paradigm in which volume consumed served as the dependent variable.35 Harris and
Booth36 also reported that a preference for salted
cereal over plain cereal was found in infants at
approximately the same age. We have hypothesized
that this developmental change in response to salt
represents, in part at least, postnatal maturation of
the ability to taste salt, an interpretation consistent
with a substantial body of evidence from animal
models.37-38
Even if there is a postnatal maturation of salt taste
perception and preference, experience with salt
could modulate this preference in infants just as it
does with adults. Other investigators36'39 have reported a correlation between one measure of exposure to salted food and relative preference for saltiness in that food. However, these data are not
conclusive as they are based on very few infants. In
our recent studies,26 no significant correlations between a measure of salt exposure and preference
have been found. Further longitudinal studies on the
development and early modification of salt taste
preference are needed. In particular, the issue of the
relation between level of exposure to salt and salt
preference should be explored at several ages in
order to understand the origin of the very high salt
preferences we have documented in some young
children.
Sodium depletion early in life may have profound
effects on later salt preference. A body of evidence,
primarily derived from studies with rats, now strongly
indicates that the salt taste system is plastic during its
development. Salt taste perception, as measured
electrophysiologically,37'38 pharmacologically,40 and
behaviorally,41-42 matures postnatally. Early restric-
1-180
Supplement I
Hypertension
Vol 17, No 1, January 1991
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tion of dietary sodium alters subsequent response to
NaCl.43 Finally, an episode of sodium depletion induced over a period of 1 or several days by a very high
dose of diuretic combined with salt-free diets produces
permanent elevation in salt intake weeks or months
after recovery, this effect appears especially robust if
the depletion occurs during early development.44
The mechanisms responsible for these effects likely
have both peripheral and central components.
Changes in the number or activity of amiloridesensitive sodium channels could account for alteration in peripheral activity of salt-sensitive neurons.
The persistent elevation of salt appetite after extreme sodium depletion, however, seems more likely
to be of central origin, possibly a consequence of the
elevation of mineralocorticoids and angiotensin that
follows sodium depletion.45-46
An analysis of available clinical studies suggests
that early sodium depletion may have profound effects on subsequent salt appetite in humans as well.
We47 searched the English language literature for
evidence that sodium depletion (occurring in clinical
entities such as Bartter's syndrome and Addison's
disease) produces a craving for salt that might be
homologous with salt appetite induced in some experimental animals after severe depletion. Surprisingly, little evidence for a human salt appetite was
found, and when it was, the onset was almost invariably in childhood. The case described by Wilkins and
Richter of a 3Vi-year-old boy with an obsessive salt
appetite is the most famous, but several other similar
cases of early childhood onset of salt appetite have
also been reported.47 The absence of persuasive
evidence for an adult onset of salt appetite suggests
that, in humans too, early depletion may have especially potent effects on later salt taste perception
and preference. This is an area ripe for further
research.48
In conclusion, humans prefer to consume substantially more salt than is necessary for the maintenance
of normal physiology. In some instances, primarily in
the salt-sensitive hypertensive individual, this may
lead to illness, and in other instances, it may contribute to the worsening of an existing illness (i.e.,
congestive heart failure, hepatic cirrhosis with fluid
retention, and increased hypertension for those on
antihypertensive medication). Some sensory and behavioral factors responsible for this high consumption have been identified, but a full explanation of
the mechanisms is lacking. To the extent that one
understands the determinants of salt intake, one may
be better able to provide guidance for successful
reduction of dietary salt intake where this is medically indicated.
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KEY WORDS • salt • diet
High salt intake. Sensory and behavioral factors.
G K Beauchamp and K Engelman
Hypertension. 1991;17:I176
doi: 10.1161/01.HYP.17.1_Suppl.I176
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Copyright © 1991 American Heart Association, Inc. All rights reserved.
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