Sensory and receptor responses to umami: an overview of pioneering
work1–3
Gary K Beauchamp
ABSTRACT
This article provides a selective overview of the early studies of
umami taste and outlines significant questions for further research.
Umami compounds such as the amino acid glutamate [often in the
form of the sodium salt monosodium glutamate (MSG)] and the nu-
cleotide monophosphates 5#-inosinate and 5#-guanylate occur nat-
urally in, and provide flavor for, many foods and cuisines around the
world. Early researchers in the United States found that the flavor of
pure MSG was difficult to describe. But they all agreed that, al-
though humans found umami compounds, when tasted alone, to be
unpalatable, subjects reported that these compounds improved the
taste of foods. This taste “dichotomy” may be partly unlearned
because it is also observed in very young infants. The uniqueness
of umami perception is based on several lines of evidence. First,
numerous perceptual studies have shown that the sensation aroused
by MSG is distinct from that of the other 4 taste qualities. Second,
biochemical studies that show the synergy of the binding of MSG
and 5#-guanylate to tongue taste tissue mirror this hallmark percep-
tual effect. Third, several specific receptors that may mediate umami
taste have recently been identified. There remain, however, a number
of puzzles surrounding the umami concept, including the molecular
basis for an apparent tactile component to umami perception, the
reason for the unpalatability of pure umami, and the functional
significance for human health and nutrition of umami detection.
Future work aimed at understanding these and other open issues
will profitably engage scientists in umami research well into the
next century. Am J Clin Nutr 2009;90(suppl):723S–7S.
INTRODUCTION
The taste properties of the sodium salt of the amino acid L-
glutamate were first described by Kikunae Ikeda, who isolated it
from seaweed in 1908 (1). He suggested that the taste was dif-
ferent from any of the consensus basic tastes (sweet, sour, salty,
bitter) and thus that it was a fifth basic taste. He further argued
that it served to allow organisms to detect and recognize sources
of protein in a manner analogous to that for sweet taste, which
may signal carbohydrate calorie sources.
In the 100 y that followed this discovery, a large body of
research has appeared that deals with the flavor (umami) of
monosodium glutamate (MSG) and the synergizing ribonu-
cleotides, inosine and guanosine monophosphates [5#-inosinate
(IMP) and 5#-guanylate (GMP)]. Although we now have a much
greater appreciation of the sensory properties of these sub-
stances, their usefulness in the flavoring of foods, and their
mechanism of action, many questions remain. Below I provide
Am J Clin Nutr 2009;90(suppl):723S–7S. Printed in USA. Ó 2009 American Society for Nutrition
a selective overview of some of the work on umami taste that
has been conducted since Ikeda’s discovery, with special at-
tention given to the work conducted outside of Japan and work
that focuses on sensory and hedonic properties. I also touch on
mechanistic studies and their potential functional significance.
Finally, several questions are proposed that require additional
investigation.
PERCEPTION
One of the early scientific articles in English on MSG, pub-
lished in 1929, described glutamic acid as a rare chemical that
imparted a meatlike taste, although at very low concentrations it
evoked a sweet taste (2). Subsequently, as the use of MSG in
foods in the United States became more common, a number of
articles appeared that discussed the sensory properties of MSG,
sometimes substantially disagreeing on the basic qualitative
properties. Indeed, there were suggestions that MSG itself had no
taste but functioned instead to enhance the other flavors in foods.
Two important early compendia on glutamate taste, published
as the proceedings of 2 symposia on MSG flavor and accept-
ability in 1948 and 1955, were held under the auspices of the US
Army Quartermaster Food and Container Institute. The military
was interested in improving the quality of the food prepared
for the troops. These symposia brought together scientists and
manufacturers to discuss and debate, among other issues, the
sensory properties of MSG. As is often the case with earlier
literature, many of the individual presentations were short on data
and long on speculation. But they did have one virtue often
missing from more recent, formalistic presentations: they en-
couraged exploration of the phenomenology of the sensory ex-
perience of MSG, alone and in food.
It is useful to quote some of the descriptions of the taste of pure
MSG by sensory specialists from the 1948 symposium:
“A desirable meat-like taste” (3)
1
From the Monell Chemical Senses Center, Philadelphia, PA.
2
Presented at the “100th Anniversary Symposium of Umami Discovery:
The Roles of Glutamate in Taste, Gastrointestinal Function, Metabolism, and
Physiology,” held in Tokyo, Japan, 10–13 September 2008.
3
Address correspondence to GK Beauchamp, Monell Chemical Senses
Center, 3500 Market Street, Philadelphia, PA 19104. E-mail: beauchamp@
monell.org.
First published online July 1, 2009; doi: 10.3945/ajcn.2009.27462E.
723S
724S BEAUCHAMP
“Its taste has all 4 components: sweetness, sourness, saltiness and
bitterness. In addition, glutamate has the capacity of stimulat-
ing the feeling nerves of mouth and throat to produce the sen-
sation known as ‘satisfaction’.” (4)
“In any event it is proper to consider glutamate as a stimulator of
the sense of feeling as well as that of taste, and the indications
are that it is unique in this capacity.” (4)
“It seems that glutamate is successful not because it adds taste,
but because it adds feeling.” (4)
“The principal effect on food flavor was a balancing, blending
and rounding out of total flavor.” (5)
“It is a sort of overall taste, flavor and feel.”(6)
“Glutamate creates a persistent, lingering flavor reaction.” (7)
A fundamental disagreement among these early investigators,
and one that still reverberates today, concerned whether the taste
properties of MSG (ie, the perception of umami) were unique, as
Ikeda had suggested, or whether they could be mimicked by an
appropriate combination of the other 4 basic taste qualities (eg,
reference 8), as Von Skramlik (9) might have proposed. The
results of such a metameric matching study (see references 10
and 11) necessary to test this hypothesis have never been pub-
lished. Moreover, a failure to find the appropriate combinations
of salt, sour, sweet, and bitter stimuli to duplicate exactly the
taste of MSG would be uninformative because one cannot prove
the negative.
As an alternative to metameric matching (and its difficulties)
for testing for unique sensory properties of umami, investigators
have resorted to using similarity scaling methods to determine
whether MSG taste differs from the other basic tastes. In an
influential study, Yamaguchi (12) used multidimensional scaling
techniques to compare the perceived flavor similarities of MSG
with sucrose, NaCl, tartaric acid, and quinine sulfate, as well as
dyads, triads, and combinations of all 4 and all 5 compounds. She
also used multiple concentrations. Very simply, she found that
within the 3-dimensional space mathematically generated from
the perceptual similarity data, MSG stood outside the space that
encompassed the 4 other taste qualities. She concluded from this
result that umami is not composed of the 4 basic tastes. Parallel
studies using similar techniques have been conducted by using
data from neural recordings (eg, reference 13) and have produced
similar results.
One problem in interpreting these data are that they assume
that MSG has only taste properties. Yet if we go back to the
discussions of the overall sensory properties of MSG in earlier
publications (summarized in the 2 Quartermaster symposia
reports), it is evident that, in addition to having a classic taste,
many investigators report that MSG also has an apparent tactile or
feeling component often described as “fullness.” This obviously
could complicate interpretation of conclusions derived from
multidimensional scaling. Could the uniqueness of the taste
properties of MSG be due to the fact that, unlike other taste
substances at the concentrations used, it alone has prominent
tactile properties? Although this is unlikely, it cannot be rejected
without further testing.
One of the most interesting characteristics of umami taste is
the synergism between MSG and the 5#-ribonucleotides IMP and
GMP. It remains controversial whether IMP and GMP them-
selves have an umami taste. When one of these substances is
placed in the mouth, a very clear umami taste can be recognized.
But it has been argued that the taste of GMP and IMP results
from synergism with the very small concentrations of free glu-
tamate present in saliva (12). This hypothesis is hard to test
critically, and unfortunately, recent molecular receptor studies in
vitro cannot resolve the issue.
In summary, early work in the United States confirmed Ikeda’s
report that MSG had a specific, perhaps unique, taste. Yet there
remained (and still remains) a curious difficulty in actually de-
fining what that taste, or more broadly, what that flavor actually
is. It would seem that if, as much subsequent molecular work
supports, umami is a distinct taste property, it is unlike the more
widely accepted 4 basic taste qualities in that it is less distinct and
more subtle.
HEDONICS
There is universal agreement that MSG, when added to many
(but not all) foods, enhances the palatability of those foods. As
early as 1948 it was recognized that this enhanced palatability
was particularly evident in soups, but was also present in a wide
variety of other nonsweet foods. It was further reported that MSG
did not enhance the acceptability of fruits, fruit juices, or some
dairy products (eg, reference 5). Palatability studies in different
cultures and across age groups have confirmed that umami
substances added to many foods increase liking (eg, reference
14).
What was not clear was the underlying mechanism by which
MSG enhanced palatability. One curious finding is that the taste
of pure MSG is generally not judged to be pleasant (eg, reference
15) unlike the taste of sugar and, in some cases, salt. This finding
in humans contrasts with work in rodents: these animals show
a clear preference for MSG and nucleotide monophosphates at
appropriate concentrations (16). This species dichotomy is an-
other way in which umami seems to differ from the other basic
tastes; we do not understand why this is the case.
A fundamental question is the extent to which the palatability-
enhancing properties of umami are learned. Human milk is quite
high in free glutamate, in contrast with cow milk, and it is thus
possible that exposure to glutamate during nursing could in-
fluence later acceptance and liking (17). Steiner (18) addressed
this question by testing newborn human infants. He gave the
infants MSG either alone or in a soup broth and then evaluated
their facial expressions in response to brief oral exposures. MSG
alone did not elicit positive facial expressions (relative to water),
although MSG in soup elicited more positive facial expressions
than soup without MSG. Steiner concluded from his results that
there is an innate component to umami liking.
We conducted studies in Mexican infants who were either well
nourished or protein-calorie malnourished to test the hypothesis
that nutritional state would feed back on taste preferences. In
particular, we wondered whether malnourished infants would
exhibit heightened preferences for MSG and/or hydrolyzed ca-
sein (which contains free glutamate). Testing consisted of giving
infants brief (1 min) access to flavor substances in baby bottles
and monitoring the amount that they consumed, a technique that
has been used successfully in many studies of infant flavor
perception and preference. In pilot work (Vazquez, Pearson, and
Beauchamp, unpublished results, 1979), we also found that
water solutions of both MSG and casein hydrolysate were not
preferred by infants; indeed, they were rejected relative to the
 PIONEERING WORK IN UMAMI TASTE
water diluent. The remainder of our work used a soup broth
similar to that used by Steiner. We found that whereas nutritional
state did affect casein hydrolysate intake (malnourished infants
preferred soup plus casein hydrolysate, whereas well-nourished
infants responded in the opposite manner), it had no effect on
MSG preference. Both malnourished and well-nourished infants
preferred (ie, consumed more in brief intake tests) MSG-con-
taining soup to soup containing no MSG (19, 20). This finding
thus fails to support the proposition that MSG signals the
presence of protein.
Our data are consistent with Steiner’s suggestion that, added to
certain foods, MSG enhances palatability even in very young
infants (the youngest in our studies were 2 mo of age). They are
also consistent with the adult judgment that MSG alone is not
palatable. But it is interesting to speculate from our results that
the sensory properties of glutamate may not actually signal the
presence of protein, as Ikeda suggested, unless they are ac-
companied by other nutrients (other amino acids) and/or flavors
that reinforce the message that nutrients are present. Indeed,
many of the natural foods high in free glutamate (eg, seaweed,
tomatoes) are not rich sources of protein. Moreover, there is very
little human evidence that protein deficiency enhances the intake
of umami-tasting foods. Our study of malnourished infants
supports this view and is one of the few human studies to date that
have examined this idea. One of the other studies (21) reported
that both elderly individuals and individuals of lower nutritional
status had a preference for higher concentrations of MSG in soup
than did individuals of higher nutritional status (as would be
predicted by this hypothesis). However, this effect could be the
result of sensory loss rather than hedonic judgment. Moreover, in
a large series of studies with protein-restricted rats, Torii et al (22)
found the reverse: protein-deficient rats avoid umami substances.
The idea that umami taste in foods signals the presence of protein
thus requires much more supporting evidence before it can be
accepted.
MECHANISM
Breakthroughs in the elucidation of receptor mechanisms
underlying taste perception are relatively recent: the first taste
receptors were identified in 1999 (16). Subsequently, an un-
derstanding of the nature of taste receptors for sweet and bitter
began to emerge, although those for sour and salty remain
something of a mystery. Umami taste reception also remains
a puzzle in part because multiple receptors have been implicated
(16).
Biochemical studies of amino acid receptors provided earlier
groundwork that led to the recent set of molecular breakthroughs.
Before the now widespread use of molecular techniques, a bio-
chemical strategy to receptor identification was used. But the
weak binding of sweet or even bitter stimuli in mammalian
preparations made this approach problematic because it was
impossible to isolate cells containing specific binding sites from
those that did not. Hence, the isolation of an enriched preparation
of cells containing taste receptor–ligand complexes could not be
achieved. This realization led investigators, among them Robert
Cagan and Joseph Brand, to select the channel catfish as a model,
on the basis of its exquisite sensitivity in identifying amino acids
such as L-arginine. This work culminated in taste receptor
identification (23). Further, it stimulated interest in the study of
725S
MSG and ribonucleotide binding in cow taste-receptor tissue, in
particular because of the high sensitivity of the synergistic effect
of added ribonucleotides. In 1984 Torii and Cagan (24) pub-
lished an important article that showed at the biochemical level
a possible mechanism underlying this perceptual synergism.
They found that binding of glutamate to bovine taste-receptor
tissue was remarkably enhanced when very small amounts of
GMP were added to the glutamate. On the basis of the analysis
of their data, they hypothesized that GMP acted to enhance the
number of sites available for MSG binding.
FUNCTION
As noted above, Ikeda’s teleological suggestion that umami
taste provides the organism with a means to detect amino acids
and protein is difficult to test and has some logical difficulties.
Nevertheless, this has remained the major hypothetical reason for
the existence of umami as a basic taste.
In 1987 Fujita (25) published a thought-provoking article
noting the anatomical similarity of endocrine cells in the gut and
taste cells in the mouth. He suggested that there might be
commonality of function. The more recent discovery of so-called
taste receptor molecular elements in gut tissue has spurred an
interest in the functional significance of this suggestion (26, 27).
Umami receptors in the mouth may be just the first step in the path
through the alimentary tract along which nutrients such as amino
acids are “recognized.” Future work in this area promises to be
very exciting.
OUTSTANDING QUESTIONS
Many unknowns, as indicated in the foregoing discussion,
remain in our understanding of the mechanisms and functions of
umami taste. A partial list of currently unanswered questions
includes the following:
1) How does glutamate provide an apparent tactile compo-
nent to foods? Does it actually stimulate tactile as well as
taste receptors, as Crocker (4) originally suggested? Could
these 2 actions be separated?
2) How is it that glutamate requires other food components to
be “appreciated”? Is this a cognitive effect (responses of
newborn infants in Steiner’s studies would argue against
this idea) or does it have a physiologic basis?
3) Multiple taste receptors have now been identified for umami
taste (23). This surfeit requires explanation: Why are mul-
tiple receptors needed? Also a puzzle are studies demon-
strating umami “blindness,” ie, individuals who apparently
cannot detect umami taste except at very high concentra-
tions (28). How could this occur if, as suggested above,
glutamate stimulates multiple taste receptors and perhaps
tactile receptors?
4) What are the physiologic roles of umami receptors
throughout the entire ingestive and digestive system?
5) Although not discussed here, several recent studies have
suggested that the intake of glutamate may have a role in
obesity that is either protective or stimulatory (29, 30). Is
there anything special about umami taste or umami sub-
stances in obesity? Also, what roles might umami substan-
ces play in the nutritive care of the sick and the elderly?
726S BEAUCHAMP
(Other articles in this supplement to the Journal include ref-
erences 17 and 31–58.)
I thank Paul Breslin, Joe Brand, and Julie Mennella for helpful comments
on earlier versions of this manuscript.
The author’s travel expenses to participate in the symposium and an
honorarium, which were paid to the author’s employer, were paid for by
the International Glutamate Technical Committee (the meeting sponsor), a
nongovernmental organization funded by industrial producers and users of
glutamate in food. The author declared no conflicts of interest.
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