Practicalities From CulinologyR How Umami Can Cont

Food and Health
Ana San Gabriel

Tia M. Rains
Gary Beauchamp Editors
Umami
Taste for Health
Food and Health
Series Editors
Jonathan Deutsch, Drexel University
Philadelphia, PA, USA
Brandy-Joe Milliron, Nutrition Sciences Department
Drexel University
The goal of this series is to provide coverage of emerging topics in food and health,
using an interdisciplinary approach that considers health not only in a functional
and human sense, but also in terms of external factors such as the environment.
Titles in the series will address growing concerns about the future health,
sustainability and quality of the food supply, as well as diet, and provide a home for
books focusing on social and environmental concerns related to food.
Ana San Gabriel • Tia M. Rains
Gary Beauchamp
Editors
Umami
Taste for Health
Editors
Ana San Gabriel Tia M. Rains
Global Communications Research and Development
Ajinomoto Co., Inc. Ajinomoto Health & Nutrition N. America
Tokyo, Japan Itasca, IL, USA
Gary Beauchamp
Monell Chemical Senses Center
ISSN 2509-6389 ISSN 2509-6397 (electronic)

Food and Health
ISBN 978-3-031-32691-2 ISBN 978-3-031-32692-9 (eBook)
https://doi.org/10.1007/978-3-031-32692-9
© The Editor(s) (if applicable) and The Author(s), 2024. This book is an open access publication.
Open Access This book is licensed under the terms of the Creative Commons Attribution 4.0
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adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit
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The editors dedicate this book to the memory
of Dr. Kunio Torii (1946–2023). During an
illustrious career, Dr. Torii made major
contributions to our understanding of the
functions and mechanisms of umami. His
professional life was characterized by his
passion for rigorous science, support and
encouragement of younger colleagues, and
dedication to enhancing human health. We
will miss his enthusiasm for work, family,
and life.
Preface
In October 1985, the first International Symposium on Umami was held in Hawaii.
The purpose of that conference was to explore physiological aspects of umami sub-
stances (mainly monosodium glutamate [MSG] and 5′-ribonucleotides) and present
findings on the physiological mechanisms of umami taste perception. Two years
after this symposium, the book Umami: A Basic Taste was published, comprising 28
chapters reflecting the presentations at this meeting. It was at this meeting and after
publication of its proceedings that the idea of umami as a novel (fifth) taste quality
became widely discussed and debated internationally in the sensory and nutrition
fields. Since this time there has been a growing scientific interest in umami, as evi-
denced by several additional symposiums, many scientific publications, and several
popular science books.
There has been a substantial amount of progress over the intervening years in
understanding of the mechanisms and functions of umami perception and prefer-
ence. It is not the purpose of this volume to review all of this work. Instead, the
current volume specifically focuses on: (1) providing summaries and analyses on
current knowledge of the perception, physiology, and molecular biology of umami;
(2) focusing on specific health-related aspects of umami; and (3) providing an over-
view of some of the culinary issues relevant to umami.
Specifically, Chap. 1 briefly discusses some of the current issues surrounding the
concept of umami from a perceptual and evolutionary perspective. Chapters 2 and 3
are devoted to basic biology of umami. These two chapters overlap to some degree,
but they provide distinctly different approaches and different emphases. This is in
part because the authors of Chap. 2 are Japanese, and it was in Japan that umami
was first identified. Japanese scientists, including these authors, have played major
roles in the scientific understanding of umami. Chapter 3 is authored by US-based
researchers who have also significantly advanced our knowledge of umami. Taken
together, these two chapters complement each other to provide a comprehensive
overview of our current understanding of the basic biology of umami.
Chapters 4–8 directly address important aspects of umami in human nutrition
and health. These chapters are authored by scientists who have played significant
investigative roles in their topics. Amid concerns about the obesity epidemic, there
vii
viii Preface
is a strong interest in understanding factors controlling satiety and satiation. It is

widely believed that umami sensation signals the presence of protein. Based on the
widespread conviction that protein is highly satiating, it has been natural to suggest
that umami may thus play an important role in satiation. Chapter 4 addresses this
issue comprehensively. Chapter 5 describes the early childhood development of
sensitivity and preference for umami stimuli. This chapter highlights how very
young infants are exposed to different levels of free (unbound) glutamate from a
very early age and how this impacts later preferences for similar flavors.
Complementing Chap. 4, it also addresses the role of umami stimuli in regulation of
infant satiety and growth.
At the other end of the age spectrum, Chap. 6 addresses umami and aging in
humans. It focuses on special nutritional challenges among the elderly population
and how umami can be used to improve their health. This topic is particularly rele-
vant given the growing proportion of elderly people in many countries around the
world. Chapter 7 highlights a particular diet-related health challenge: the necessity
to reduce consumption of excess salt (sodium) in virtually every country around the
world. Umami and salt are closely allied (both NaCl and MSG contain sodium), and
now considerable research indicates that umami stimuli can replace a certain amount
of salt in many foods, thereby reducing an individual’s overall sodium consumption.
Chapter 7 comprehensively reviews the growing literature on this topic. Chapter 8
provides a broader overview of health and umami, presenting umami as a seasoning
that can increase the appeal of foods that align with human and planetary health.
While there are several barriers to consuming healthful diets, taste is among the
most important. Global data analysis has shown that poor diets lead to chronic dis-
eases. Chapter 8 considers how the use of umami, due to its taste and physiological
functions, could enhance acceptance of foods that may reduce the risk of chronic
diseases and their contributions to premature mortality. To conclude our volume,
Chap. 9 reminds us that umami is, after all, a food-related concept and that culinary
usage of umami ingredients has a long history in many cuisines worldwide. This
chapter also includes practical guidance for umami usage from experts in the field.
A note on the book cover: Green is evocative of health, plants, and growth, and
peas are one of many fresh vegetables high in glutamate.
Tokyo, Japan Ana San Gabriel

Itasca, IL, USA Tia M. Rains
Philadelphia, PA, USA Gary Beauchamp
Acknowledgements
This book would not have been possible without the support of the International
Glutamate Information Service (IGS) and the Monell Chemical Senses Center. We
also thank Kumiko Ninomiya and Patricia J. Watson for their valuable editorial
assistance.
ix
Contents
1
Introduction: Umami as a Taste Percept �� 1
Gary Beauchamp
2 Umami and MSG �� 7
Ryusuke Yoshida and Yuzo Ninomiya
3
Umami Taste Signaling from the Taste Bud to Cortex �� 43
Eugene R. Delay and Stephen D. Roper
4
Umami and Salty: A Cooperative Pair�� 73
Aubrey Dunteman and Soo-Yeun Lee
5 Umami and Satiety�� 101
Martin R. Yeomans
6 Umami Taste: Inborn and Experiential Effects on Taste
Acceptance and Satiation During Infancy�� 127
Ana San Gabriel and Julie A. Mennella
7 Umami and Healthy Aging�� 147
Minoru Kouzuki and Katsuya Urakami
8
Umami Taste as a Component of Healthy Diets�� 165
Ana San Gabriel and Tia M. Rains
9 Practicalities from Culinology®: How Umami Can Contribute
to Culinary Arts and Sciences�� 183
Chris Koetke, Lauren Miller, and Jonathan Deutsch
Index�� 197
xi
Contributors
Contributors
Gary Beauchamp Monell Chemical Senses Center, Philadelphia, PA, USA

Eugene R. Delay Regis University, Denver, CO, USA
Emeritus Faculty at the University of Vermont in Burlington, Burlington, VT, USA
Jonathan Deutsch Drexel University, Philadelphia, PA, USA
Aubrey Dunteman University of Illinois, Urbana, IL, USA
Ana San Gabriel Global Communications, Ajinomoto Co., Inc., Tokyo, Japan
Chris Koetke Ajinomoto Health & Nutrition North America, Itasca, IL, USA
Minoru Kouzuki Tottori University, Tottori, Japan
Soo-Yeun Lee University of Illinois, Urbana, IL, USA
Julie A. Mennella Monell Chemical Senses Center, Philadelphia, PA, USA
Lauren Miller Drexel University, Philadelphia, PA, USA
Yuzo Ninomiya Kyushu University, Fukuoka, Japan
Tia M. Rains Research and Development, Ajinomoto Health & Nutrition North
America, Itasca, IL, USA
Stephen D. Roper Department of Physiology and Biophysics and the Department
of Otolaryngology, Miller School of Medicine, University of Miami, Coral
Gables, FL, USA
Katsuya Urakami Tottori University, Tottori, Japan
Martin R. Yeomans University of Sussex, Brighton, UK
Ryusuke Yoshida Okayama University, Okayama, Japan
xiii
Chapter 1
Introduction: Umami as a Taste Percept
Gary Beauchamp
Umami is now commonly identified as the fifth “basic” taste quality, joining sweet,
salty, sour, and bitter. It must be emphasized that the term taste quality refers to
human (and perhaps other species) psychological representations, not to the ligands
that elicit those representations; for example, NaCl elicits a salty taste in humans,
but sodium chloride itself is not “salty.” The four traditional basic taste qualities
have a very long and deep history in human experience. As detailed in a recent
review (Beauchamp, 2019), sweet, salty, sour, and bitter, along with pungency and
astringency (these last two are tactile qualities, not taste qualities), were identified
as the basic building blocks of perceived taste in Chinese, Indian, and Greek writ-
ings dating back several thousands of years. Moreover, these four taste qualities, as
well as the two tactile qualities, also make up much of what is reported to be the
current taste world in many independent, relatively isolated cultures around the
world (Beauchamp, 2019).
These four basic taste qualities likely exist to provide vital information on the
health and safety of potential foods. Sweet and salty substances are generally highly
palatable, signaling vital nutrients: calories and sodium. Bitter compounds, with
generally negative hedonic qualities, usually signal danger or poison. However, bit-
ter has also been seen as a signal for medicinal value, both historically and currently,
in many cultures around the world (Beauchamp, 2019). Functionally, sour remains
a puzzle. Among several hypotheses to account for it, one is that it acts as an inhibi-
tory signal for unripe fruit that could also injure the oral cavity, and another is that
it could signal the presence of certain micronutrients (Breslin, 2019; Liman &
Kinnamon, 2021). It is significant that, for sweet, salty, and bitter, the perceptual
signal is generally identical with the actual function of the signaling molecules.
That is, most sweet compounds in nature are calorie-rich, virtually all salty
G. Beauchamp (*)
Research and Development, Monell Chemical Senses Center, Philadelphia, PA, USA
e-mail: [email protected]
© The Author(s), 2024 1

A. San Gabriel et al. (eds.), Umami, Food and Health,
https://doi.org/10.1007/978-3-031-32692-9_1
2 G. Beauchamp
substances in nature contain sodium, and most molecules that are bitter act as poi-
sons at least in high concentrations.
At the beginning of the twentieth century, the Japanese chemist Kikunae Ikeda
identified the glutamate ion as inducing a novel taste quality that he proposed was a
signal for protein, analogous to sweetness being a signal for energy or calories (see
Chaps. 2 and 3). He described this novel quality as “the peculiar taste we feel as
umai [meaning, according to the translators, meaty, brothy, or savory].” He called
this taste umami. Subsequent studies further identified the ribonucleotides
5′-inosinate and 5′-guanylate as synergistic enhancers of this novel taste, although
for humans and some other species, they may not produce a taste on their own.
Beginning in the late 1960s and 1970s, additional researchers also proposed umami
as a novel fifth basic taste. This idea gained momentum from a scientific meeting
held in 1985 and the publication of the proceedings Umami: A Basic Taste
(Kawamura & Kare, 1987). The evidence used to support the proposal that umami
was the fifth basic taste at that time consisted primarily of human sensory studies
and animal model studies of behavior and physiology.
The discovery in the 1990s and 2000s of specific taste receptors responsive to
amino acids (see Chaps. 2 and 3 in this volume) gave substantial impetus to the idea
that umami might be the fifth basic taste. Yet even the proponents of umami as a
basic taste acknowledge that it differs significantly from the classic four (e.g.,
Hartley et al., 2019): it is more subtle, which may account for it not being identified
historically or in most traditional cultures, and unlike the other appetitive taste qual-
ities, sweet and salty, relatively pure solutions of monosodium glutamate (MSG) are
very rare in nature and are not palatable for human adults or infants (see Chap. 2).
Moreover, unlike sweet, salty, and bitter, the perceptual quality of umami for
humans does not directly signal the presence of the purported nutrient, proteins
(which generally have no taste), or even amino acids. Indeed, many umami-rich
foods are not naturally high in protein (Breslin, 2013).
Although much current research on umami focuses on its apparently unique taste
to humans, early descriptions of the umami percept, or perceptual characteristics,
included a strong component that is best described as tactile rather than taste
(Beauchamp, 2009). Whether this tactile percept, sometimes called mouthfeel,
results from a true ability of MSG to engage somatosensory rather than taste path-
ways, or whether it is mediated by anatomically defined taste pathways, as has
recently been argued by Yamamoto and Inui-Yamamoto (2023), remains to be deter-
mined. Nevertheless, mouthfeel is an extremely important and salient component of
the umami percept, and it merits future research. For example, does this attribute
have any causal relationship to the observation that umami sensations are involved
in satiety and satiation?
1 Introduction: Umami as a Taste Percept 3
1.1 Species Differences in Umami Perception
Do other species detect umami? Taste qualities are human-derived percepts, so it

can be problematic to discuss umami in the context of other species—which of
course is also true of the other basic taste qualities. The words salty, sweet, bitter,
and sour refer to sensory properties that humans perceive. However, for these taste
qualities, there are at least two reasons to believe that at least some other animal
species perceive something similar to what humans perceive. First, many of the
compounds we classify as sweet or salty and as bitter or sour elicit similar behaviors
in several other well-studied species. For example, rats and mice tend to avoid bitter
compounds and are attracted to many compounds humans describe as sweet—sweet
is good and bitter is bad, for the most part. What is more, if a rodent is made ill by
exposure to a simple sugar such as glucose (to which a tasteless purgative, e.g., has
been associated), it will not only avoid this taste when presented again but also
avoid many other simple sugars and even some nonnutritive compounds humans
describe as sweet, such as saccharin. This suggests that all these compounds elicit a
common percept—a common taste quality.
Is the umami taste similar across species? The apparent answer is no. Umami
stimuli for humans are restricted to MSG and, to a lesser extent, aspartate and are
synergistically enhanced by some ribonucleotides. However, this specificity is not
evident for rodents (Nelson et al., 2002) and perhaps many other species. In some
species, MSG appears to be sweet or salty, whereas in others, glutamate may elicit
the same percept (neither sweet nor salty) as many other amino acids. That is, the
putative receptor for umami in humans (the dimer T1R1/T1R3; see Chaps. 2 and 3)
may be quite different in other species, due to molecular changes in its binding
affinities and more central projections and thus different in the perceptual character-
istics it elicits in many other species. Consequently, it is much more problematic to
speak of the “umami receptor” in species other than humans than it is to speak of the
“sweet receptor” or the “bitter receptors” in some other species.
1.2 Umami Perception in Humans
Why, from a functional and evolutionary perspective, is human perception of umami

elicited almost exclusively by glutamate and, to a lesser degree, by aspartate?
Several suggestions have been proposed to explain this—while none are definitive,
each may have merit. Following the lead of Ikeda (2002) who was writing in 1909,
Breslin (2013, 2019) has suggested that humans developed a preference (and pre-
sumably a specific receptor) for glutamate and ribonucleotides as markers for pro-
tein. But many high-protein foods do not have a strong umami taste. Breslin’s idea
4 G. Beauchamp
is that the specific umami taste quality is particularly human because it signals eas-
ily digested protein as formed during cooking (see Chaps. 5 and 9) and fermenta-
tion. He noted that fermented foods also have the nutritional advantage of providing
easy access not only to amino acids but also to probiotic bacteria. Thus, he partially
attributes specificity of human umami perception and preference for food manipula-
tion by our human ancestors. As Breslin (2019, p. 15) summarizes: “We can pre-
sume that this taste, which we call savory or umami, was initially related to
fermentation.” Although this explanation has merit, it fails to explain why other
primate species as well as nonprimate mammals (which do not cook or ferment their
foods) also appear to have a receptor system focused on glutamate and/or
ribonucleotides.
A second recent approach toward understanding the human specificity of umami
has been suggested in a comprehensive evaluation of the T1R1/T1R3 receptor in 17
species of primates (Toda et al., 2021). Each of these primate species has a func-
tional T1R1/T1R3, but this receptor varies in which stimuli engage it most effec-
tively. To evaluate these differences, Toda and colleagues tested the receptor
responses to glutamate and to ribonucleotides 5′-inosinate and 5′-guanylate sepa-
rately in each of these species and attempted to associate relative responsiveness to
these compounds to the primate’s dietary habits. They concluded that for primates
that consume primarily insects, this receptor is specialized for detection of unbound
(free) ribonucleotides, which is consistent with the presence of large amounts of
these molecules in insects. In contrast, unlike in humans, the T1R1/T1R3 receptor
in these species does not respond well to glutamate. They proposed that ribonucleo-
tide sensitivity was the ancestral response of all primates. Subsequently, the T1R1/
T1R3 receptor evolved to respond specifically to glutamate in a variety of primate
species, including human precursors. These species consume primarily leaves,
which are low in free ribonucleotides but are relatively rich in free glutamate. They
concluded that glutamate sensitivity could be useful for these species for detecting
dietary protein in their plant-based diets. They further speculated that this respon-
siveness to the free glutamate in plant leaves also functions to mask or inhibit the
bitterness of leaf-based secondary metabolites, thereby heightening leaf palatabil-
ity. Although Toda and colleagues focused on primate analyses, they also investi-
gated glutamate responsiveness in the isolated T1R1/T1R3 receptor in several other
species of mammals, including mice, cats, dogs, horses, and pigs. The receptors of
all these nonprimate species except pigs were relatively unresponsive to glutamate,
but all were highly responsive to the ribonucleotides. This further emphasizes the
novel specialized nature of the umami response in humans and closely related
primates.
One additional aspect of umami in humans bears mentioning. Human milk is
particularly rich in glutamate, as are the milks of several other closely related pri-
mates (gorillas, chimpanzees, and even rhesus macaques; Rassin et al., 1978; Davis
et al., 1994; Sarwar et al., 1998). These species’ T1R1/T1R3 receptors are also
highly responsive to glutamate (Toda et al., 2021). Could there be a causative rela-
tionship between specificity for glutamate and a high concentration of glutamate in
breast milk? Thus, leaf eating may not be the only driving force toward high
1 Introduction: Umami as a Taste Percept 5
receptor specificity for free glutamate and thus for umami as a positive stimulus in
primates that are more closely related to humans.
1.3 Conclusion
Although many mysteries about umami taste remain, we nevertheless know much
more about this potent sensory percept and its health-related aspects now than we
did even 20 years ago. Taste and associated oral sensations such as mouthfeel pro-
vide the last chance for an organism to decide whether to ingest or reject a particular
food. Taste signals the potential worth and possible danger of foods. Umami percep-
tion in the oral cavity is now recognized as a significant force in nutrition and health,
as are the associated physiological effects of umami stimulations of T1R1/T1R3
and perhaps other receptors in the oral cavity and elsewhere in the body. The chap-
ters in this book dramatically illustrate this, describing what we know and calling
attention to what we still do not know about mysterious umami.
References
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work. The American Journal of Clinical Nutrition, 90(3), 723S–727S. https://doi.org/10.3945/
ajcn.2009.27462E
Beauchamp, G. K. (2019). Basic taste: a perceptual concept. Journal of Agricultural and Food
Chemistry, 67(50), 13860–13869. https://doi.org/10.1021/acs.jafc.9b03542
Breslin, P. A. (2013). An evolutionary perspective on food and human taste. Current Biology, 23,
R409–R418.
Breslin, P. A. S. (2019). Chemical senses in feeding, belonging, and surviving; or, are you
going to eat that? (Elements in Perception). Cambridge University Press. https://doi.
org/10.1017/9781108644372
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Journal of Nutrition, 124(7), 1126–1132. https://doi.org/10.1093/jn/124.7.1126
Hartley, I. E., Liem, D. G., & Keast, R. (2019). Umami as an “alimentary” taste: a new perspective
on taste classification. Nutrients, 11(1), 182. https://doi.org/10.3390/nu11010182
Ikeda, K. (2002). New seasonings. Chemical Senses, 27, 847–849.
Kawamura, Y., & Kare, M. R. (1987). Umami: a basic taste—physiology, biochemistry, nutrition,
food science. Dekker.
Liman, E. R., & Kinnamon, S. C. (2021). Sour taste: receptors, cells and circuits. Current Opinion
in Physiology, 20, 8–15.
Nelson, G., Chandrashekar, J., Hoon, M., et al. (2002). An amino-acid taste receptor. Nature, 416,
199–202. https://doi.org/10.1038/nature726
Rassin, D. K., Sturman, J. A., & Gaull, G. E. (1978). Taurine and other free amino acids in milk of
man and other mammals. Early Human Development, 2(1), 1–13.
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6 G. Beauchamp
Toda, Y., Hayakawa, T., Itoigawa, A., Kurihara, Y., Nakagita, T., Hayashi, M., Ashino, R., Melin,
A. D., Ishimaru, Y., Kawamura, S., Imai, H., & Misaka, T. (2021). Evolution of the primate glu-
tamate taste sensor from a nucleotide sensor. Current Biology, 31(20), 4641–4649.e5. https://
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Gary Beauchamp is Distinguished Member, and Director and President Emeritus, of the Monell
Chemical Senses Center. He has served as a scientific advisor to numerous governmental and pri-
vate organizations and has published widely on the senses of taste and smell. His current research
interests include the role of taste and flavor in food and beverage perception and acceptance, the
genetics of chemosensation, and the development and aging of taste and smell. He received his
Bachelor’s Degree in Biology from Carleton College and his Ph.D. in Biopsychology from the
Pritzker School of Medicine of the University of Chicago.
Open Access This chapter is licensed under the terms of the Creative Commons Attribution 4.0
adaptation, distribution and reproduction in any medium or format, as long as you give appropriate
credit to the original author(s) and the source, provide a link to the Creative Commons license and
indicate if changes were made.
The images or other third party material in this chapter are included in the chapter’s Creative
Commons license, unless indicated otherwise in a credit line to the material. If material is not
included in the chapter’s Creative Commons license and your intended use is not permitted by
statutory regulation or exceeds the permitted use, you will need to obtain permission directly from
the copyright holder.
Chapter 2
Umami and MSG
Ryusuke Yoshida and Yuzo Ninomiya
2.1 Historical Context of Umami and MSG
2.1.1 Discovery of Umami Taste by Kikunae Ikeda
The sense of taste, which is elicited by chemical compounds in the oral cavity, plays
a critical role for food intake. When we consume sugar, table salt, vinegar, or coffee,
we clearly feel a sweet, salty, sour, or bitter taste, respectively. In addition to these
four basic tastes, umami is now considered the fifth basic taste. Umami taste was
first described about 110 years ago (Ikeda, 1908, 1909, 2002; Lindemann et al.,
2002) by Kikunae Ikeda (1864–1936; see Fig. 2.1). Ikeda, a chemistry professor at
the Imperial University of Tokyo, had studied in Germany for 2 years in the labora-
tory of Friedrich Wilhelm Ostwald at the University of Leipzig. At that time, four
tastes (sweet, sour, salty, and bitter) were considered “pure” tastes, whereas others,
such as hot, metallic, alkaline, and astringent tastes, were not considered “pure”
tastes, because chemical compounds eliciting these sensations were detected, at
least in part, by the somatosensory system rather than by the taste system.
Ikeda had been interested in the taste of the Japanese seaweed broth dashi
because he believed that dashi clearly contained another (pure) taste, which was
different from sweet, salty, sour, and bitter tastes and was also recognized in meat
and fish dishes. He intended to isolate the principal taste substance from the sea-
weed Laminaria japonica, the main ingredient for dashi. After conducting many
procedures, such as aqueous extraction, removal of large-scale contaminants
R. Yoshida (*)
Okayama University, Okayama, Japan
Y. Ninomiya
Kyushu University, Fukuoka, Japan

https://doi.org/10.1007/978-3-031-32692-9_2
8 R. Yoshida and Y. Ninomiya
Fig. 2.1 Dr. Kikunae

Ikeda (photo taken in
1923). (Image from the
Umami Information Center
website, https://www.
umamiinfo.jp/what/
whatisumami/)
(mannitol, sodium, and potassium chloride), and lead precipitation, he finally

obtained pure crystals of a single substance that he identified as glutamic acid. He
proposed to call the taste of glutamic acid umami, a word derived from the Japanese
adjective umai (delicious). Indeed, he noted that the taste of glutamic acid crystals
was perceived as umami taste after its sour taste had faded and that the salts of glu-
tamic acid (sodium, barium, calcium, and potassium) had strong umami taste. The
term umami as a taste was first mentioned in his original Japanese paper, but in later
publications, he used the English phrase glutamic taste as a scientific term repre-
senting the peculiar taste of glutamate (glutamic acid) that is different from all other
well-defined taste qualities (Ikeda, 1912).
Ikeda described several aspects of the taste intensity of glutamate (Ikeda, 1909,
2002). He reported that the taste recognition threshold for monosodium L-glutamate
(MSG) was about 1/3000 (1.6 mM), which is lower than that of sucrose (1/200,
15 mM) and NaCl (1/400, 43 mM). Although the taste intensity of glutamate
increased as its concentration increased, changes in the taste intensity of increasing
concentrations of glutamate were likely to be smaller than those of sweet, salty,
sour, and bitter tastes—umami taste did not become extremely strong even at high
glutamate concentrations. Ikeda also described the taste of mixtures. For example,
2 Umami and MSG 9
the taste of glutamate was substantially decreased by addition of acids. This may be
due to the addition of hydrogen ions to the glutamate solution, yielding a no dissoci-
ated form of glutamate (hydrogen glutamate), leading to decreased concentrations
of the glutamic acid anion, the taste stimulus for umami taste. Mixing salt (NaCl)
with a glutamate solution increased the palatability of ionic glutamic acid, although
a weak salty taste did not enhance the intensity of the glutamate taste. The sweet-
ness of sugars was not affected by the taste of ionic glutamic acids, but the taste of
ionic glutamic acid was decreased by strong sweetness. In addition, the taste of
sweet stimuli and the taste of ionic glutamic acid had some similarities: some peo-
ple perceived the taste of ionic glutamic acid as sweet at a concentration close to the
threshold.
Ikeda also addressed the stereochemical structure of the amino acid associated
with umami taste (Ikeda, 1909, 2002), but at that time it was difficult to explain the
relationship between molecular structure and taste. He further considered umami
taste from the viewpoint of its nutritional value. Because meat extract contains a
certain amount of glutamic acid, along with other amino acids, he reasoned that the
taste of glutamate could be an indicator of the presence of nutritive foods, particu-
larly of protein. Therefore, a preference for umami taste may have evolved to
encourage intake of such protein-rich foods. Although Ikeda discussed preference
for umami taste, he noted in a later publication that umami taste (glutamic taste) by
itself was not palatable or delicious (Ikeda, 1912), and this has also been noted by
others (Yamaguchi, 1991; Halpern, 2002). When MSG is added to the appropriate
foods, it increases the palatability of those foods (Halpern, 2000). Therefore, in
Europe and America, umami tastants have often been regarded as flavor enhancers
or potentiators. In summary, the basic logic and characteristics of umami taste were
described in the first paper on the taste of glutamate by Ikeda (1909). His work
formed the foundation of studies on umami taste.
It was subsequently noted that some nucleotides have taste characteristics similar
to glutamate (umami). Shintaro Kodama, a pupil of Ikeda, isolated 5′-inosinic acids
from another ingredient of dashi, katsuobushi (dried skipjack, bonito flakes), as a
constituent having a taste similar to that of glutamate (Kodama, 1913). About a half
century after Ikeda’s work, Akira Kuninaka found 5′-guanylic acid from dried black
mushrooms (shiitake, Lentinus edodes) as another umami tastant. He also found
that the taste intensity of umami was greatly enhanced by mixing of MSG and
5′-ribonucleotides, the phenomenon known as umami synergism (Kuninaka, 1960).
Synergism between glutamate and nucleotides, discussed in more detail below, is a
hallmark of umami taste and is widely used in cooking to enhance the palatability
of foods. Such synergism has been reported in physiological and psychological
studies. For example, gustatory nerve responses to MSG were greatly enhanced by
adding inosine 5′-monophosphate (IMP) or guanosine 5′-monophosphate (GMP) in
mice (Ninomiya & Funakoshi, 1987, 1989a), dogs (Kumazawa & Kurihara, 1990),
and rats (Yamamoto et al., 1991). The first biochemical data indicating synergistic
effects of glutamate and nucleotides were demonstrated in bovine taste papillae
(Torii & Cagan, 1980). More recently, the molecular mechanism underlying umami
synergism has been elucidated (Zhang et al., 2008).
2.1.2 First Symposium on MSG by the US Army
MSG had been used in food industries, restaurants, and some home consumers to
improve palatability in the United States since the 1930s. From the 1920s, the
Japanese Imperial Army had tackled methods to improve the quality of army rations,
including the use of MSG to improve the taste of rations such as canned foods.
During World War II, the US Army employed MSG to improve the quality of foods
for troops. After the war, in 1948 and 1955, the US Army Quartermaster Food and
Container Institute held two symposia on MSG flavor and acceptability. At these
symposia, scientists and manufacturers discussed and debated various aspects of
MSG, including its production, its use as a flavoring agent, and its sensory proper-
ties (Quartermaster Food and Container Institute, 1948; Research and Development
Associates 1955; Yamaguchi & Ninomiya, 1998; Beauchamp, 2009).
The usefulness of MSG in recipes of the US Army’s master menu was thor-
oughly explored. In one study, preference tests of 50 foods and recipes were con-
ducted with ~2150 individuals for 18 months (Girardot & Peryam, 1954). Among
the 50 foods and recipes, addition of MSG clearly improved the palatability of 25
foods and recipes and weakly enhanced that of 3 foods and recipes. In contrast, 4
foods and recipes were worsened and 18 were not affected by adding MSG. Overall,
the palatability of meat, fish, and vegetable dishes tended to be greatly improved by
the addition of MSG, whereas that of cereals, milk products, and sweet dishes was
not. Thus, MSG has the potential to enhance the palatability of some but not
all foods.
Regarding sensory properties of MSG, neither the concept of umami nor the
synergistic effect of MSG and nucleotides had been established at that time.
Therefore, how its taste was represented by sensory specialists is worth noting to
understand the history of umami taste. From Yamaguchi and Ninomiya (1998),
some descriptions for MSG taste at that time were as follows (italics indicate the
points by the authors):
• Taste of MSG had a tingling feeling factor, and persistency of taste sensation
presented in the whole of the mouth region, including the roof of the mouth and
the throat. It was hard to describe the sensation other than to call it a feeling of
satisfaction. These suggest that MSG stimulated nerve endings lying within the
buccal cavity and stimulated the sense of feeling as well as that of taste
(Crocker, 1948).
• 0.1–0.3% of MSG had a sweet saline taste accompanied by some astringency. It
stimulated all surfaces of the tongue and oral cavity, producing a slight sensation
of furriness on the tongue and a mild but lasting aftertaste (Cairncross, 1948).
• When a small amount of MSG was placed on the tongue, salivary secretion was
increased and lasted for approximately half an hour. It produced a slight sensa-
tion of furriness on the tongue and mild stimulation in the throat and the back
part of mouth. There was a sensation of bloom, i.e., the taste seemed to spread
rapidly inside of mouth and had an after effect on the tongue (Cairncross &
Sjöström, 1948).
2 Umami and MSG 11
• MSG had an effect of aroma, without contributing any noticeable odor itself. The
principal effect on food flavor was regarded as balancing, blending, and rounding
out the total flavor without contributing any noticeable odor or taste, except that
it was very noticeable in certain fruits and dairy products. MSG enhanced mouth-
fullness and satisfaction (Cairncross, 1948).
• Glutamic taste was not unique and could be duplicated by a mixture of the four
tastes (Crocker & Henderson, 1932).
Compared to four basic taste qualities (sweet, salty, sour, and bitter), such repre-
sentations of taste of MSG were complicated and diverse and elicited disagreement.
The apparent taste of glutamate was likely to include tactile, olfactory, visceral, and
other sensations. At that time (and to some degree even now), the “taste” of gluta-
mate elicited controversy, but the effects of glutamate on oral sensations, such as
“tingling,” “persistency,” “satisfaction,” “mouthfullness,” and “aftertaste,” were
noted, suggesting that glutamate may stimulate something other than or in addition
to taste in the oral cavity (see Sect. 2.4). In some cases, sweet and salty tastes were
mentioned as the taste of MSG. This may be attributed, at least in part, to the sodium
component of MSG, since low concentrations of NaCl were recognized as sweet
when subjects were adapted to water (Bartoshuk, 1974).
2.1.3 Chinese Restaurant Syndrome and MSG Safety
From the 1930s to the 1960s, production and consumption of MSG became preva-
lent worldwide. Then, in 1968, a letter to the editor titled “Chinese-Restaurant
Syndrome” by Robert Ho Man Kwok, MD, was published in the New England
Journal of Medicine (Kwok, 1968). He reported that he had experienced a strange
syndrome after he had eaten foods in a Chinese restaurant, with symptoms of numb-
ness, general weakness, and palpitations. One of the causes of these symptoms, he
speculated, was the high sodium content of the Chinese foods, which may produce
hypernatremia, leading to intracellular hypokalemia, causing such symptoms.
Because MSG seasoning contains the sodium ion and was used to a great extent in
Chinese dishes, he hypothesized that MSG may be a cause of such symptoms. This
letter in the New England Journal of Medicine elicited a large reaction (Schaumburg,
1968; McCaghren, 1968; Menken, 1968; Rose, 1968; Rath, 1968; Beron, 1968;
Kandall, 1968; Gordon, 1968, Davies, 1968).
This original letter, as well as many comments about it often supporting the
symptoms listed, led investigators to experimentally test MSG as the culprit.
Schaumburg et al. (1969) reported that intake of MSG produced such typical symp-
toms as burning sensations, facial pressure, and chest pain in all but one test subject.
Morselli and Garattini (1970) carried out a study on 24 healthy volunteers using a
double-blind technique and showed no significant differences in symptoms between
intake of MSG and placebo. But Himms-Hagen (1970) criticized their results as
they did not use susceptible subjects. At that time, Olney (1969) reported that sub-
cutaneous injections of MSG (0.5–4 mg/g body weight) in 2- to 9-day-old mice
caused extensive damage to neurons in the hypothalamus and other areas of the
brain. A similar result was obtained in one infant rhesus monkey (Macaca mulatta)
(Olney & Sharpe, 1969). Olney and Ho (1970) also reported that orally adminis-
trated MSG (and aspartate and cystatin) induced hypothalamic damage in
infant mice.
Such reports had great impact on the general public, and “Chinese restaurant
syndrome” (or MSG toxicity) became widely known. However, many following
studies showed little or no relationship between MSG intake and the typical symp-
toms described for Chinese restaurant syndrome (Freeman, 2006; Greisingera et al.,
2016). Kenny and Tidball (1972) explored the human reactions to oral MSG and
confirmed the results of Morselli and Garattini (1970). Kerr et al. (1977, 1979)
investigated aversive symptoms associated with foods and found no respondent who
met the criteria for all three aversive symptoms for MSG (tightness and burning
sensation in the head and chest, numbness). In 1986 the FDA’s Advisory Committee
on Hypersensitivity to Food Constituents concluded that MSG posed no threat to
the general public, and in 1987 the Food and Agriculture Organization of the United
Nations (FAO)-World Health Organization (WHO) Joint Expert Committee on
Food Additives placed MSG in the safest category of food ingredient (Tracy, 2016).
Figure 2.2 gives a timeline of umami discovery, use, and research.
Indeed, ingested glutamate (and glutamate produced by degradation of proteins in
the intestine) in ordinary foods is used for oxidative fuel and as a precursor for other
amino acids, glutathione, and N-acetyl glutamate (Blachier et al., 2009; Burrin &
Stall, 2009). In healthy human volunteers, jejunal and ileal L-glutamate content is
greatly increased at 3 hours after the ingestion of a test meal, but the concentration of
glutamate in venous blood plasma was only slightly increased at 1 h after the inges-
tion of a test meal (Adibi & Mercer, 1973). In addition, MSG ingestion with a meal
in healthy human subjects did not result in any significant increase in plasma gluta-
mate level 15–360 min after ingestion (Ghezzi et al., 1985). Experiments in the piglet
using a newly developed labeled tracer demonstrated that >95% of enteral glutamate
but only 5% of the enteral glucose was utilized by the mucosa (Reed et al., 2000).
Although experimental conditions were different, these studies suggest that only a
small amount of glutamate is taken into the blood through the intestine and that most
glutamate, when taken with food, is used as fuel and as resources for bioactive sub-
stances in the gastrointestinal tract after absorption of glutamate in the intestine.
Regarding effects of MSG intake on the brain, administered MSG in animals did
not significantly affect brain glutamate levels in infant or adult animals (Airoldi
et al., 1979; Garattini, 1979, 2000). Furthermore, extracellular glutamate in the
hypothalamus or striatum of rats was not increased when MSG was administrated
as a component of food (Bogdanov & Wurtman 1994; Monno et al., 1995). These
data suggest that brain glutamate levels are not greatly increased when MSG is
ingested along with meals. Although neurotoxic effects of glutamate are well known
(Lau & Tymianski, 2010), the conclusion was that normal intake of MSG (with
foods) does not damage the brain. However, the impression of MSG as a food addi-
tive and also the impression of glutamic taste became worse in the 1960s and 1970s;
such an impression still remains in some people today (Yeung, 2020).
2 Umami and MSG 13
1900
1908: Ikeda’s Japanese patent

1909: First Japanese paper on umami by Ikeda
1913: Finding of umami taste of 5’-inosinic acid
1920 1920s: Use of MSG in army rations by Japanese Imperial Army
1930s: Prevalence of consumption of MSG in USA
1940
1948: First symposium on monosodium glutamate in Chicago, USA
1955: Second symposium on monosodium glutamate in Chicago, USA
1960 1960: Finding of umami taste of 5’-guanylic acid

1960: Finding of umami synergism
1968: Chinese restaurant syndrome
1978: First international conference on glutamic acid in Milan, Italy

1980 1982: Foundation of The Umami Research Organization
1985: International symposium on umami in Hawaii, USA
1986: Safety of MSG by FDA
1987: Safety of MSG by FAO-WHO
1990: Umami symposium in Sicily, Italy
1996: Finding of umami receptor (taste-mGluR4)
1998: Umami symposium in Bergamo, Italy
2000
2002: Finding of umami receptor (TAS1R1/TAS1R3)
2008: Umami symposium in Tokyo
2020
Fig. 2.2 The chronology of umami taste and monosodium L-glutamate

2.1.4 Umami as a Basic Taste
In 1978, the first international conference on glutamic acid (“The International

Symposium on Biochemistry and Physiology of Glutamic Acid”) was held in Milan,
Italy (see Fig. 2.2). This symposium focused on such topics as the sensory and
dietary aspects of glutamate, metabolism of glutamate, roles of glutamate in the
central nervous system, and evaluation of the safety of glutamate (Filer Jr. et al.,
1979). These researchers did not describe glutamate as umami or list its taste as one
of the basic tastes, but the term umami began to spread among international scien-
tists during this period. In 1982, researchers in fields of physiology, biochemistry,
nutrition, and food science established the Umami Research Organization study
group to promote research on umami. This organization held the first international
symposium on umami in Hawaii (1985). The purpose of this symposium was to
explore physiological aspects of the effects of umami substances on flavor evalua-
tion of foods and beverages and to present research findings on the physiological
mechanisms of umami taste perception (Kawamura & Kare, 1987). The proceed-
ings of this symposium, “Umami: A Basic Taste,” provided a comprehensive view
of umami studies, including general concepts, developmental aspects, receptor
mechanisms, psychometric analyses, physiology and behavior, brain mechanisms,
and nutrition and behavior, as all of these topics relate to umami taste (Fig. 2.3).
This symposium drew international participants and contributors, including investi-
gators from Japan, the United States, England, France, Switzerland, Israel, and
Mexico. This symposium established the term umami internationally. Now we use
umami as a scientific term representing taste of glutamate (and also nucleotides).
Subsequently, this organization held umami symposia in Sicily (1990), Bergamo
(1998), and Tokyo (2008) and held sessions on umami taste in International
Symposium on Olfaction and Taste (ISOT) meetings in Sapporo (1993), San Diego
(1997), Kyoto (2004), San Francisco (2008), Stockholm (2012), Yokohama (2016),
and Portland (2020).
When searching the keyword “umami” in PubMed, we find a few articles from
the 1980s. The number of articles per year in the 1980s and 1990s was less than 10,
except for 1991 (21 reports, containing the proceedings of second international
umami symposium in Sicily, held in 1990) and 1999 (12 reports). After that, the
number of articles per year rapidly increased, reaching 176 in 2020. During this
period, the most pivotal study on umami taste was the identification of umami taste
receptors (Lindemann et al., 2002). The first report demonstrating the receptor for
glutamate in peripheral taste tissue was published in 1996 (Chaudhari et al., 1996).
This study demonstrated that a taste-specific variant of metabotropic glutamate
receptor 4 (taste-mGluR4), lacking most of the N-terminal extracellular domain,
was expressed in taste buds of rats. In 2002, another G-protein-coupled receptor,
the TAS1R1 + TAS1R3 heterodimer, was reported to function as an umami (amino
acid) receptor (Li et al., 2002, Nelson et al., 2002). Furthermore, the variant of
metabotropic glutamate receptor 1 was reported to be expressed in taste tissue and
may function as an umami taste receptor (San Gabriel et al., 2005). The findings of
2 Umami and MSG 15
Fig. 2.3 Cover of the proceedings for the 1987 symposium “Umami: A Basic Taste”
specific receptors for glutamate (and other amino acids) in taste cells emphasized
that umami taste is different from sweet, salty, sour, and bitter taste. Besides previ-
ous evidence of physiological and psychological studies on umami taste (described
in Sect. 2.2), these molecular studies supported the concept that umami is one of
the basic tastes. More than a century after the discovery of umami by Ikeda, umami
has become well accepted internationally in the scientific field of taste perception.
2.2 Umami and Other Basic Tastes
From ancient times, sensations of taste were classified, divided, or categorized into
qualities (elements). In ancient Greece, Aristotle proposed seven elements of taste
(flavor): sweet, bitter, salty, sour, pungent, astringent, and rough. In Ikeda’s first
report on umami, he noted as follows:
In the past it was said that there are five taste qualities: sour, sweet, salty, bitter, and hot. A
hot sensation is just a skin mechanical sensation; therefore, today’s scientists do not regard
this sensation as taste. Furthermore, such qualities of metallic, alkaline and astringent are
not considered to be tastes, because they cannot be separated from the sensation accompa-
nied by tissue damage. Therefore, physiologists and psychologists recognize only the four
tastes sour, sweet, salty and bitter. (Ikeda, 1909, 2002)
Thus, for thousands of years in writings by Chinese and Indian scholars, as well as
in traditional medicinal practices around the world, sour, sweet, salty, and bitter
have been accepted as distinct, primary, or basic taste qualities (Beauchamp, 2019).
Each of these tastes is considered to provide an organism with specific information
about energy sources (sweet), minerals (salty), acids (sour), and poisonous com-
pounds (bitter) in foods and drinks. Typical taste compounds used in taste researches
are sucrose (sweet), NaCl (salty), citric acid (sour), and quinine (bitter). To consider
whether umami is a basic taste or not, some definitions of a basic taste are required.
There have been many attempts to identify appropriate criteria for defining a basic
taste (Beauchamp, 2019). One of the most widely accepted set of criteria was pro-
posed by Kurihara in the proceedings of symposium “Umami: A Basic Taste.” He
proposed that a basic taste could be defined as follows (Kurihara, 1987, 2015):
1. A basic taste should be found universally in many foods.
2. A basic taste should not be produced by any combination of other basic tastes.
3. A basic taste should be independent of other basic tastes as proven by psycho-
physical and electrophysiological studies.
4. A specific receptor for a basic taste should exist.
Since umami taste fulfills these definitions, umami can be considered a fifth
basic taste.
2.2.1 Umami Substances in Foods
Compounds eliciting sweet, bitter, sour, and salty tastes are found naturally in many
foods at detectable concentrations. Do umami compounds also exist naturally in
many foods? Indeed, glutamate and umami ribonucleotides are widely distributed in
natural foods (Ninomiya, 1998a, 2002; Yoshida, 1998). Glutamic acid is a promi-
nent component in such foods as meats, fishes, and vegetables (Table 2.1). Some
vegetables and seafood contain considerable amounts of free glutamate (Table 2.2).
It is noteworthy that human milk contains a considerable amount of free glutamate
2 Umami and MSG 17
Table 2.1 Amino acid composition of selected foods

Amount (mg/100 g)
Amino acid Beef Chicken Tuna Oyster Tomato Potato Cow’s milk Human milk
Ala 840 1600 1400 360 19 45 100 36
Arg 900 1700 1400 340 19 74 100 32
Asp 1300 2200 2400 570 71 320 250 86
Cys 160 250 256 81 8.9 20 29 24
Glu 2100 3400 3500 840 240 260 620 170
Gly 730 2100 1100 360 18 44 59 22
His 500 910 2400 130 12 26 88 26
Ile 630 980 1200 220 15 50 170 51
Leu 1100 1700 2000 370 25 78 310 99
Lys 1200 1900 2300 400 25 82 260 66
Met 360 600 760 140 6.3 24 83 15
Phe 570 900 970 220 18 59 150 42
Pro 610 1400 850 290 17 56 300 92
Ser 540 950 950 250 22 54 150 41
Thr 620 1000 1100 260 17 54 130 43
Trp 160 240 300 58 5 17 41 15
Tyr 480 750 860 180 14 36 120 40
Val 700 1100 1300 250 17 79 210 56
Data extracted from Standard Tables of Food Composition in Japan (MEXT, 2015)
Table 2.2 Free glutamate in selected foods
Food product Free glutamate (mg/100 g)

Beef 0.56–19.1
Pork 6.0–18.5
Chicken 7.1–13.0
Tuna 3–5
Salmon 6.2–25.4
Oyster 123–207
Sea urchin 67–219
Tomato 93.6
Potato 90.6
Cow’s milk 3
Human milk 18.3
Cheese 41.2–453
Soy sauce 782
Cured ham 636
Data extracted from Database for Free Amino Acid Compositions of Foods (Japan Society of
Nutrition and Food Science, 2013)
(18.3 mg/100 g). It is possible that exposure to glutamate during nursing could
influence later acceptance and liking (Mennella et al., 2009).
The amount of free glutamate is increased in fermented and processed foods
such as cheese, soy sauce, and cured ham. In general, content of free glutamate in
foods is increased by storage, maturation, ripening, cooking, and other processing.
In the case of meats, free glutamate increases during storage. Free glutamate in
tomato increases as the fruit matures: fully ripe tomatoes contain ten times the con-
centration of free glutamate as green tomatoes. The content of free glutamate in
cheeses and cured ham also increases during their ripening. Thus, glutamate can be
a natural stimulant (tastant) when we eat various food stuffs. Nucleotides such as
IMP and GMP are also abundant in some foods (Table 2.3). Dried skipjack (bonito)
contains a large amount of IMP, while dried black mushroom contains a large
amount of GMP. Both of these materials have been used to isolate umami com-
pounds (Kodama, 1913; Kuninaka, 1960). Therefore, nucleotides also are natural
tastants in many foods. Taken together, the umami compounds glutamate and nucle-
otides are abundant in many foods and act as stimulants to taste organs, fulfilling
one of criteria for a basic taste.
Although some foods contain glutamate abundantly, the contribution of gluta-
mate (and also other substances) to the taste of foods needs to be investigated. To do
this, omission tests have been conducted (Fuke & Konosu, 1991). In these tests, first
the chemical composition of a food is analyzed and determined. Then, the mixture
of pure chemicals representing chemical compounds of the food is made, and the
taste is tested to determine whether the synthetic mixture has a taste similar to the
Table 2.3 5′-Ribonucleotides in foods

Amount (mg/100 g)
Food product 5′-Inosinic acid 5′-Guanylic acid 5′-Adenylic acid
Beef 70.7 3.7 7.5
Pork 200.2 2.2 8.6
Chicken 201.3 5.3 13.1
Tuna 286 — 5.9
Snow club 5.0 4.0 32.0
Prawn — / 86.8
Scallop — — 172.0
Sea urchin — — 28.0
Dried skipjack 474 / 52
Asparagus — — 4.0
Tomato — — 20.8
Potato (raw) — — /
Potato (boiled) — 2.3 3.8
Black mushroom (raw) — — /
Black mushroom (dried) — 150 /
Data from Ninomiya (1998a)
— not detected, / not analyzed
2 Umami and MSG 19
Table 2.4 Extractive components in the leg meat of snow crab
Component mg/100 g Component mg/100 g Component mg/100 g

Ala 187 Tyr 19 Trimethylamine oxide 338
Arg 579 Val 30 Glucose 17
Asp 10 Adenine 1 Ribose 4
Glu 19 Adenosine 26 Lactic acid 100
Gly 623 Betaine 357 Succinic acid 9
His 8 Cytosine 1 ADP 7
Ile 29 Guanine 1 AMP 32
Leu 30 Homarine 63 CMP 6
Lys 25 Hypoxanthine 7 GMP 4
Met 19 Inosine 13 IMP 5
Phe 17 Ornithine 1 Cl– 336
Pro 327 Sarcosine 77 K+ 197
Ser 14 Taurine 243 Na+ 191
Thr 14 -Methylhistidine 3 PO43– 217
Trp 10 -Aminobutyric acid 2
Data from Fuke and Konosu (1991)
Abbreviations: ADP adenosine 5′-diphosphate, AMP adenosine 5′-monophosphate (5′-adenylic
acid), CMP cytidine 5′-monophosphate, GMP guanosine 5′-monophosphate, IMP inosine
5′-monophosphate. Compounds whose omission changed the taste of crab leg in omission tests
are shaded
original food. After that, one or more of the compounds are omitted from the syn-
thetic mixture, and the taste of the mixture is tested to determine whether the omit-
ted mixture still tastes similar to the original food. If the omission of a certain
compound changes the taste, that compound may be essential for the taste of the
original food.
Using this procedure, essential taste compounds for boiled snow crab meat were
analyzed. The extracts from the leg meat of boiled snow crab contained many com-
pounds, including glutamic acid (Table 2.4). Among these compounds, omission of
glutamate, glycine, arginine, adenosine 5′-monophosphate (AMP), GMP, and
sodium and chloride ions changed the taste of crab meat. Glutamate and 5′-ribonu-
cleotides were particularly important in increasing the overall identity and prefer-
ence. Similar to the crab meat, the taste of other seafood such as abalone, sea urchin,
scallop, short-necked clam, dried skipjack, and salted salmon eggs was unfavorably
altered by omission of glutamate or 5′-ribonucleotides. Thus, umami compounds
are essential components for the taste of many types of seafood.
2.2.2 Interaction Between Umami and Other Tastes
Do umami substances such as glutamate and nucleotides affect other basic tastes
and vice versa? As indicated earlier (see Sect. 2.1), Ikeda first noted that (a) the taste
of glutamate was substantially decreased by the addition of acids, (b) a weak salty
taste did not enhance the intensity of glutamate taste, and (c) the sweetness of sugars
was not affected by the taste of ionic glutamic acids, whereas strong sweetness
weakened the taste of ionic glutamic acid (Ikeda, 1909, 2002). To reveal the interac-
tion between umami and other basic tastes, many psychophysical studies have since
been carried out, with somewhat inconsistent results. For example, Lockhart and
Gainer (1950) reported that MSG did not affect the thresholds of sugar and salt solu-
tions. Mosel and Kantrowitz (1952) reported that administration of MSG reduced
the threshold of sour and bitter tastes but not of sweet and salty tastes. Van Cott et al.
(1954) demonstrated that MSG at a concentration 0.75 times threshold reduced the
threshold of sweet and salty tastes but not of bitter and sour tastes. To clarify the
effect of umami substances on the taste of sweet, salty, sour, and bitter, Yamaguchi
and Kimizuka (1979) measured the thresholds of four basic tastants (sucrose, NaCl,
tartaric acid, quinine) with or without 5 mM MSG or IMP. The detection threshold
for quinine sulfate was slightly increased by addition of IMP but not MSG. The
threshold of tartaric acid was considerably raised by adding MSG or IMP, but those
of sucrose and NaCl were not affected by addition of MSG or IMP. The effect of
IMP on bitter thresholds may be explained by a masking effect by the slight bitter
side taste of IMP, and that of MSG and IMP on sour taste may be caused by
changes in pH.
Conversely, effects of other tastants on the detection threshold of MSG were
investigated (Yamaguchi, 1987). In this case, the threshold of MSG was not greatly
increased by addition of other tastants, even at high concentrations, except for
higher concentrations of sucrose. From these results, there may be some interac-
tions between umami and other tastes, but these interactions may be explained by
physicochemical properties or side tastes of umami substances. Thus, umami taste
is likely to be independent from other basic tastes.
2.2.3 Psychophysical and Multidimensional Studies

of Umami Independence
Although Ikeda described umami (glutamic taste) as distinct from sweet, salty, sour,
and bitter tastes, many US researchers believed that it could be duplicated by a mix-
ture of the four basic tastes. For example, Crocker and Henderson (1932) reported
that the taste of MSG could be duplicated by mixing sucrose, NaCl, tartaric acid,
and caffeine. The taste of glutamate is generally weak, and the addition of MSG to
an appropriate food increases the flavor, pleasantness, and acceptability of the food
(Halpern, 2000). Therefore, glutamate has been often considered to be a flavor
enhancer rather than a taste substance itself.
In 1916, the German psychologist Hans Henning proposed the concept of the
taste tetrahedron (Henning, 1916, 1984). If each of basic tastes is arranged at one of
the apices of a tetrahedron, the taste of a certain compound will be represented as a
point within that tetrahedron. This idea is essentially that taste perception of any
compound or mixture of compounds could be duplicated by mixtures of four pri-
mary tastes (sweet, salty, sour, and bitter). This implies that if a certain taste could
not be depicted within the taste tetrahedron, that taste should be categorized as a
specific, primary, or basic taste.
2 Umami and MSG 21
This concept has been adopted to show the independence or distinctiveness of

umami taste. By mathematical analysis of psychological and physiological data
using a method called multidimensional scaling (MDS), tastes of many substances
can be represented within three-dimensional space. MDS can thus provide a visual
representation of the pattern of proximities among a set of objects. Using MDS of
human psychophysical data, the similarity of the tastes of amino acids, including
MSG, was analyzed (Yoshida & Saito, 1969). This report included an MDS three-
dimensional representation of taste of amino acids, NaCl, and MSG, at 12 times the
concentration of their thresholds. The taste tetrahedron had apices of salty (NaCl),
bitter (tryptophan, etc.), sour (glutamic acid, aspartic acid), and sweet (alanine, gly-
cine); MSG was found to be positioned outside of the tetrahedron. However, this
report did not demonstrate a clear segregation of the taste of MSG.
Schiffman et al. (1980) used MDS to show the similarity of the taste of sodium
salts, including MSG, in humans. They used 13 sodium salts, as well as sucrose
(sweet), citric acid (sour), and quinine (bitter). In their MDS representation, the
taste tetrahedron has four vertices (sucrose, citric acid, quinine, and NaCl), and the
position of MSG was separate from these tastes, outside of the taste tetrahedron.
Furthermore, Yamaguchi (1987) used MDS to examine similarities of 21 taste stim-
uli of single and mixture solutions of sucrose (sweet), NaCl (salty), tartaric acid
(sour), quinine sulfate (bitter), and MSG. In the three-dimensional representation of
the results, the four basic tastes were located at the four vertices of the tetrahedron
(Fig. 2.4, dashed lines). All mixtures of four basic tastes were located on the edges,
the faces, the inside, or the vicinity of the tetrahedron. In contrast, MSG was clearly
positioned at a distance from the tetrahedron. These mathematical analyses of
human psychological data on taste similarity suggest that the taste of MSG is not
composed of the four basic tastes and has characteristics different from those of the
four basic tastes, fulfilling one of the criteria for a basic taste. However, it should be
noted that this could be caused by the presence of other, nontaste sensory properties
of MSG, such as tactile sensations.
MDS was also used to analyze taste response properties in experimental animals.
Ninomiya and Funakoshi (1987, 1989a) investigated responses in mice of gustatory
nerve fibers in the chorda tympani nerve (innervating the anterior part of the tongue)
and the glossopharyngeal nerve (innervating posterior part of the tongue). They
found multiple fibers showing responses to MSG and also synergism between MSG
and GMP. In the glossopharyngeal nerve, they identified MSG-best fibers that did
not show responses to sweet, salty, sour, or bitter tastants. This provided strong
evidence for the existence of a neural pathway that specifically sends umami infor-
mation to the brain. The MDS of these responses demonstrated that umami com-
pounds (MSG, GMP, IMP, MSG+GMP) formed a cluster present outside of the
tetrahedron circumscribed by salty (NaCl), sour (HCl), bitter (quinine), and sweet
(sucrose, fructose, maltose, glucose, saccharin) tastes, especially when using the
data of glossopharyngeal nerve fibers or of all of tested fibers.
Ninomiya and Funakoshi (1987, 1989b) also investigated taste similarity of 16
test stimuli in mice by using a conditioned taste aversion paradigm. If mice were
conditioned to avoid either MSG, monosodium L-aspartate (MSA), disodium
Fig. 2.4 Multidimensional scaling produced this three-dimensional representation of taste simi-
larities among 21 taste stimuli (individually and as mixtures): S, sucrose, sweet; N, NaCl, salty; T,
tartaric acid, sour; Q, quinine, bitter; and M, monosodium L-glutamate, umami. SNTQ1–5 consist
of different concentrations of S, N, T, and Q. X1 (X2, X3), dimension 1 (2, 3); (+), positive value.
The dashed lines outline the classic taste tetrahedron with salty, sweet, sour, and bitter at the verti-
ces. (Image from the Umami Information Center website, https://www.umamiinfo.jp/what/attrac-
tion/taste, modified from Yamaguchi (1987))
5′-inosinate (IMP), or disodium 5′-guanylate (GMP) alone, they also avoided the
other three compounds as well. This phenomenon is called generalization. Such
data indicate taste similarity among MSG, MSA, IMP, and GMP in mice. The MDS
of these data demonstrated that a cluster of umami compounds (MSG, MSA, IMP,
GMP, MSG+GMP) was outside of the taste tetrahedron apices composed of salty
(NaCl), bitter (quinine), sour (HCl), and sweet (sucrose, saccharin, glucose, fruc-
tose, glycine, L-glutamine). Thus, in mice the taste of glutamate may be perceived
as different from the other basic tastes. In investigations of the gustatory nerve fibers
of chimpanzees, similar MDS showed a separateness of umami taste from other
basic tastes (Hellekant et al., 1997a). In a hierarchical cluster analysis, they found
an M-subcluster of gustatory fibers that responded robustly to umami substances
(MSG, GMP, MSG+GMP). Using MDS, the positions of MSG, GMP, and
MSG+GMP were apart from all other tastants, suggesting that umami taste is dis-
tinct from other basic tastes in the chimpanzee at the level of the peripheral gusta-
tory nerve fibers.
Taste responses have also been investigated in the higher-order neurons and ana-
lyzed by MDS. Baylis and Rolls (1991) investigated taste responses of neurons in
the taste cortex of macaques to understand the neural encoding of glutamate taste in
a primate. Using five tastants (glucose, NaCl, HCl, quinine, and MSG), they
2 Umami and MSG 23
recorded 190 neurons and found single neurons tuned to respond best to MSG. MDS
of these data showed that MSG was located apart from the tetrahedron composed of
the other four basic tastes. In addition, Rolls et al. (1996) examined responses to the
glutamate ion and IMP in neurons of the taste cortex. MSG-best neurons responded
well to glutamic acid, and the response to glutamic acid correlated well with that to
MSG but not to glucose, NaCl, HCl, or quinine. The response to IMP also correlated
well with that to MSG. In MDS, glutamic acid was located near MSG, which was
distant from the tetrahedron composed of sweet, sour, salty, and bitter tastants.
Therefore, in the taste cortex of macaques, umami taste (MSG, IMP, and glutamic
acid) may be encoded differently from the other basic tastes. In summary, the MDS
of taste data in humans and some experimental animals strongly emphasizes the
different characteristics of umami taste compared with the other four basic tastes.
2.2.4 Umami Receptors
Marked additional evidence showing that umami is a basic taste comes from molec-
ular studies of taste receptors. The first biochemical evidence for an umami taste
receptor was demonstrated by using bovine taste papillae (Torii & Cagan, 1980). In
this study, binding of L-[3H]glutamate to bovine circumvallate papillae was mea-
sured, showing that the addition of nucleotides substantially enhanced binding of
glutamate to a preparation of bovine taste papillae, providing molecular evidence
for umami synergism. Beginning around 2000, molecular studies have led to the
identification of several receptors for the basic tastes. In 2000, G-protein-coupled
receptors named taste 2 receptors (TAS2Rs) were identified as bitter taste receptors
(Chandrashekar et al., 2000; Matsunami et al., 2000). In 2001, TAS1R3 was identi-
fied as the gene product of the Sac locus and was shown to function as a sweet
receptor together with TAS1R2 (Bachmanov et al., 2001; Kitagawa et al., 2001;
Max et al., 2001; Montmayeur et al., 2001; Nelson et al., 2001; Sainz et al., 2001).
Regarding umami taste receptors, a taste-specific variant of mGluR4 (taste-mGluR4)
was first identified as a candidate receptor for umami taste expressed in taste tissue
of rats (Chaudhari et al., 1996, 2000). Thereafter, the dimer TAS1R1 + TAS1R3 was
identified as another candidate receptor for umami (amino acid) taste (Li et al.,
2002, Nelson et al., 2002). Furthermore, taste-mGluR1 was also reported to be a
candidate receptor for umami (San Gabriel et al., 2005). Together with the salt taste
receptor ENaC (epithelial sodium channel; Chandracheker et al., 2010) and sour
taste receptor OTOP1 (otopetrin 1; Teng et al., 2019; Zhang et al., 2019), one or
more taste receptors have been identified for each of the five basic tastes. These
molecular studies suggest that umami receptors are different from receptors for
other basic tastes, providing additional evidence that umami is a distinct basic taste.
It is noteworthy that TAS1R3 is a common component both for sweet and for umami
receptors; such sharing is not found for the other classes of receptors for bitter, salty,
and sour.
2.2.5 Neural Pathways for Umami Taste
Based on the abovementioned studies, umami taste fulfills all conditions for the
definition of a basic taste as listed by Kurihara (1987, 2015): (1) found universally
in many foods, (2) not produced by any combination of other basic tastes, (3) inde-
pendent of other basic tastes by psychophysical and electrophysiological studies,
and (4) has a specific receptor. A fundamental question is how umami taste is coded
in the neural system. Is there any specific neural pathway for umami taste? As men-
tioned earlier, the existence of a specific neural pathway for umami taste was dem-
onstrated in single-fiber recordings in mice (Ninomiya & Funakoshi, 1987). In
addition, more recent studies have demonstrated the existence of umami-best (or
umami-specific) gustatory nerve fibers (Yasumatsu et al., 2012) and neurons in the
geniculate ganglion (Barretto et al., 2015; Wu et al., 2015), which contains cell bod-
ies of gustatory nerve fibers in mice. At the taste cell level, MSG-best taste cells
were found in mice in both circumvallate papillae (Maruyama et al., 2006) and
fungiform papillae (Niki et al., 2011). These taste cells may transmit their informa-
tion to MSG-best gustatory nerve fibers, forming a peripheral neural pathway con-
ducting information of umami taste to higher-order neurons.
In the brain, how basic taste qualities are represented in the primary taste cortex
of mice was examined by using an in vivo two-photon calcium imaging technique
(Chen et al., 2011). They found that each taste quality is represented in its own sepa-
rate cortical field, forming a “gustotopic” map in the insula. The umami cortical
field, which is apart from sweet, bitter, and NaCl cortical fields, was specifically
tuned to umami stimuli and contained fewer neurons responding to the other four
taste qualities. Thus, umami may be coded in such an umami cortical field in the
insula. Given that umami-best (or umami-specific) cells exist in the taste buds, the
taste ganglions, and the taste cortex, it would not be surprising if a dedicated neural
pathway coding umami taste from the peripheral to the central nervous system is
present in mice.
2.3 Differences in Umami Taste
2.3.1 Species Differences
Many animal species have been studied for their taste sensitivity to umami sub-
stances. As mentioned above, mice have some specific neural lines for umami taste
(Ninomiya & Funakoshi, 1987, 1989a). At the behavioral level, mice can discrimi-
nate the taste of MSG from that of sweet (sucrose, saccharin, fructose, glucose, and
maltose), bitter (quinine), sour (HCl), and salty (NaCl) (Ninomiya & Funakoshi,
1987, 1989b). Thus, mice have the ability to sense MSG as a taste different from
others. However, some differences in umami sensitivity might exist among mouse
strains. Many inbred mouse strains have been developed and used in various
2 Umami and MSG 25
studies. Among them, B6 strains have higher avidity for sweeteners than do 129
strains (Lush, 1989). Regarding umami taste, umami synergism between MSG and
GMP varies across strains, in the order of C3H > B6 > BALB strains, at the gusta-
tory nerve level (Ninomiya et al., 1992). B6 mice consumed more MSG than did
129 mice in behavioral tests, but gustatory nerve responses to MSG did not differ
between B6 and 129 strains (Bachmanov et al., 2001). Nonetheless, mice can sense
the taste of glutamate.
In hamsters, Yamamoto et al. (1988) examined electrophysiological and behav-
ioral responses to umami substances. Single-fiber recording of chorda tympani
nerve fibers demonstrated that some fibers responded to MSG, IMP, and/or
MSG+IMP. These responses to umami substances were highly correlated with NaCl
responses but poorly with other taste stimuli, suggesting no or little specific neural
line for umami taste in the chorda tympani nerve of hamsters. In whole-nerve
recordings of the glossopharyngeal nerve, no response was observed for 0.3 M
MSG, 0.3 M IMP, or 0.3 M NaCl. In addition, synergistic enhancement between
MSG and IMP was not observed in whole-nerve recordings of the chorda tympani
nerve. At the behavioral level, hamsters conditioned to MSG showed avoidance to
NaCl and vice versa. Thus, hamsters may not discriminate the taste of MSG and
NaCl and also may not sense synergism between glutamate and nucleotides.
In rats, single chorda tympani nerve responses to umami substances demon-
strated that some fibers responded to MSG, GMP, and MSG+GMP (Sato et al.,
1970). Among these fibers, synergism between MSG and GMP was found in
sucrose-sensitive fibers. In whole-nerve recordings, the chorda tympani nerve
showed clear synergism between MSG and IMP (Yamamoto et al., 1991). Chorda
tympani nerve responses to MSG and IMP were mostly inhibited by amiloride, an
epithelial sodium channel blocker, whereas those to MSG+IMP and monopotas-
sium L-glutamate (MPG) + IMP were suppressed by Gymnema sylvestre extract, a
sweet taste inhibitor. In behavioral experiments, rats conditioned to avoid umami
substances showed avoidance to sucrose but not to NaCl, HCl, or quinine. If rats
were conditioned to avoid sucrose, they also avoided umami substances (Yamamoto
et al., 1991). Such a link between sweet and umami substances has been reported in
other studies (Chaudhari et al., 1996; Stapleton et al., 2002; Heyer et al., 2003,
2004). These neural and behavioral data indicate that rats may have difficulty distin-
guishing between umami and sweet taste.
Umami responses have been investigated in animals other than rodents. In the
dog, neural responses to umami substances from the chorda tympani nerve showed
a large synergism between MSG and GMP or IMP in most mongrel dogs and
between MSG and GMP, IMP, or AMP in beagles (Kumazawa & Kurihara 1990).
Addition of nucleotides did not enhance responses to NaCl, sucrose, HCl, or qui-
nine, suggesting canines have an umami receptor that shows a synergistic effect
between glutamate and nucleotides. In the pig, gustatory nerve fiber responses in the
chorda tympani and glossopharyngeal nerve showed the existence of M-type fibers
with large responses to MSG (Danilova et al., 1999). M-type fibers in the glosso-
pharyngeal nerve showed high specificity to umami stimuli compared to those in the
chorda tympani nerve, suggesting that the umami information derived from the
glossopharyngeal nerve is more important than that from the chorda tympani nerve
for discriminating umami stimuli from other stimuli. In the calf, single-fiber
responses of the chorda tympani nerve demonstrated that some fibers responded to
MSG but most also showed responses to NaCl, LiCl, and urea (Hellekant et al.,
2010). In case of the calf, taste fibers dominantly responding to MSG may not exist
in the chorda tympani nerve. However, it is possible that these fibers exist in the
glossopharyngeal nerve, as is the case for pigs and mice. Because biochemical evi-
dence for a synergistic effect between glutamate and nucleotide was demonstrated
using bovine circumvallate papillae (Torii & Cagan, 1980), a receptor system under-
lying umami synergism should exist in taste cells of the posterior part of the bovine
tongue. In primates, as mentioned previously, an M-subcluster of gustatory fibers
was found in the chorda tympani nerve of chimpanzees (Hellekant et al., 1997a).
Single cortical neurons tuned to respond best to MSG were found in the taste cortex
of macaques (Baylis & Rolls, 1991). In addition, MSG-best fibers were found in the
glossopharyngeal nerve of rhesus monkeys (Hellekant et al., 1997b).
Taken together, many species of animals are sensitive to umami substances, but
species do differ in sensitivity and neural representation of umami signals. More
details on species differences of umami taste from the view of receptors are
described in Chap. 3 of this volume. In brief, genes for umami receptor components,
TAS1R1 and/or TAS1R3, are pseudogenized (inactive) in some species, including
the sea lion, the bottlenose dolphin, and the giant panda (Li et al., 2010; Jiang et al.,
2012). These animals lack functional TAS1R1 + TAS1R3 receptors and thus may
not taste umami substances.
2.3.2 Tongue Regional Differences
Sensitivity of the tongue differs by region. In experimental animals, these regional

differences are inferred by finding differences between tongue areas innervated by
different nerves. In the case of mice, regional differences of sensitivity to amiloride
were reported (Ninomiya et al., 1991, Ninomiya, 1998b). Amiloride selectively sup-
pressed NaCl responses of the chorda tympani nerve innervating the anterior part of
the tongue by about 50% of control but did not inhibit those of the glossopharyngeal
nerve innervating the posterior part of the tongue. Similarly, gurmarin, a sweet
receptor blocker for mouse and rat isolated from the plant Gymnema sylvestre,
selectively suppressed sweet responses of the chorda tympani nerve but not of the
glossopharyngeal nerve (Ninomiya et al., 1997). Umami substances such as MSG,
IMP, and MSG+IMP contain the sodium ion. Therefore, responses of the chorda
tympani nerve to these substances are partly suppressed by amiloride. In the chorda
tympani nerve, fibers showing large responses to MSG and synergism between
MSG and IMP predominantly responded to sucrose (S-best fibers). Gurmarin almost
completely suppressed responses of this type of fiber not only to sucrose but also to
MSG+IMP. However, chorda tympani nerve fibers predominantly sensitive to
umami substances (M-type fibers) did not exhibit such suppression of responses to
2 Umami and MSG 27
umami substances by gurmarin (Ninomiya et al., 2000; Yasumatsu et al., 2006). In

contrast, the glossopharyngeal nerve of mice contains a much greater number of
M-type fibers and showed greater responses to umami substances (Ninomiya et al.,
2000). In line with these data, at the behavioral level, transection (cutting) of the
glossopharyngeal nerves affected licking behavior of mice in a conditioned taste
aversion paradigm (Ninomiya & Funakoshi, 1989b). Mice conditioned to avoid
MSG showed no avoidance to sucrose, NaCl, HCl, or quinine (no generalization to
other taste stimuli), but these mice did show avoidance to NaCl (generalization to
NaCl) if the glossopharyngeal nerves but not the chorda tympani nerves were bilat-
erally transected. Thus, the glossopharyngeal nerve likely sends taste information
for umami, which can be discriminated from that of the other basic tastes in mice.
The presence of M-type fibers in the glossopharyngeal nerve was also demonstrated
in rhesus monkeys (Hellekant et al., 1997b).
Although there are no data on gustatory nerve responses to umami substances in
humans, psychophysical experiments have been done using a filter-paper test, in
which a small piece of filter paper soaked with the taste solution is applied directly
to the area of interest on the tongue. These studies demonstrated that umami sensi-
tivities stimulated with MSG, IMP, and MSG+IMP were higher on the posterior
than on the anterior part of the tongue (Yamaguchi & Ninomiya, 2000; see Fig. 2.5).
3
160mmol/L sucrose
2
Locus1 2 3 4 5
1
3
0
3
320mmol/L NaCl
2 5
2cm
1 2 4
Evaluation score
1 1cm
0 2cm
3 3
40mmol/L DL-tartaric acid 160mmol/L MSG
2 2
1 1
0 0
3 3
0.625mmol/L quinine sulfate 160mmol/L IMP
2 2
1 1
0 0
3 3
Water 40mmol/L MSG + 40mmol/L IMP
2 2
1 1
0 0
Sweet Salty Sour Bitter Umami Sweet Salty Sour Bitter Umami
Taste quality
Fig. 2.5 Evaluation scores as mean certainty ratings (0 = none or uncertain, 1 = likely, 2 = fairly,
3 = absolutely) for each taste quality perceived for each taste stimulus at five loci of the tongue.
Stimuli were taste solutions on a filter paper disk (n = 30). MSG, monosodium L-glutamate; IMP,
inosine 5′-monophosphate. (Modified from Yamaguchi and Ninomiya (2000))
Similar results were reported by other researchers (Feeney & Hayes, 2014). Based
on this research, it is suggested that the posterior part of the tongue may play the
major role in detection and discrimination of umami-specific (or umami-dominant)
information.
2.4 Distinctive Phenomena of Umami Taste
Although Ikeda laid the foundation for umami taste more than 100 years ago, sub-
sequent evidence on the taste of glutamate has solidified the concept of a unique
umami taste. However, several puzzling phenomena about umami taste remain to be
elucidated. In Western countries, the taste of glutamate has been described as
“savory,” “mouthfullness,” or “brothlike” (Ninomiya, 2002). As mentioned by sen-
sory specialists from the 1948 symposium on MSG flavor and acceptability
(described in Sect. 2.1 above), the “taste” or flavor of glutamate was described as
“tingling,” “persistent,” and “satisfying” (Beauchamp, 2009). In addition, “long-
lasting,” “aftertaste,” and “stimulation in the throat” were keywords representing
taste of glutamate in that symposium (Yamaguchi & Ninomiya, 1998). Of course,
glutamate in the oral cavity stimulates taste receptor cells on the tongue. However,
from these descriptions of its taste, glutamate may engage sensory pathways other
than those detected by the sense of taste, such as tactile (touch) sensations. For
example, the oral sensation of acids may consist of sour taste and nociceptive or
painful sensations. Although wild-type mice avoided drinking acid solutions such as
citric acid, genetic ablation of sour taste receptor OTOP1 did not affect avoidance of
acid solutions (Zhang et al., 2019). Similarly, ablation of trigeminal neurons
expressing TRPV1 (transient receptor potential member V1) did not eliminate
avoidance of acid solutions. In contrast, mice lacking both the Otop1 gene and
TRPV1-expressing trigeminal neurons showed reduced avoidance of an acid solu-
tion, suggesting both taste and nociceptive components are required for perception
and avoidance of acid stimuli. Further studies are required to elucidate whether oral
glutamate stimulates sensations other than taste.
2.4.1 Intensity of Umami Taste
As mentioned by Ikeda, one of its characteristics that distinguishes umami from

other tastes is that umami does not become extremely strong even at high concentra-
tions of glutamate. At the suprathreshold level, the relationship between the subjec-
tive taste intensity and the concentration of tastants can be expressed by the
following equations (Yamaguchi, 1998):
MSG: S = 9.69 log2(x/0.0195)
Sucrose: S = 14.98 log2(x/0.873)
NaCl: S = 15.50 log2(x/0.0943)
2 Umami and MSG 29
100 Quinine Tartaric

sulfate acid NaCl Sucrose
Intensity of taste
50
MSG
0
10-3 10-2 10-1 1 10 100
Concentration (g/100ml)
Fig. 2.6 Relationship between taste intensity and concentration. (Modified from Yamaguchi (1987))
Tartaric acid: S = 14.45 log2(x/0.00296)

Quinine sulfate: S = 14.16 log2(x/0.000169)
Here, x is the concentration of each taste stimulus (g/dl) and S is the subjective taste
intensity (the taste intensity of saturated sucrose solution is represented as S = 100).
Although the subjective taste intensity of MSG, like that of the other four basic
tastants, follows Fechner’s law, where the subjective sensation is proportional to the
logarithm of the stimulus intensity, the slope of MSG’s concentration-intensity
function is less steep than that of others (Fig. 2.6). Ikeda used an analogy to express
this characteristic: it is like the color of yellow, which does not appear to intensify
when the concentration is increased; in contrast, sweet taste is like the color red,
which does intensify as the concentration increases. The mechanisms underlying
this unique taste characteristic of umami are still unknown, but this characteristic
may prevent us from noticing umami taste in many foods—umami is much less
salient in foods than are sweet, sour, salty, and bitter.
2.4.2 Synergism
Umami synergism was first reported by Kuninaka (1960). He noticed that the
umami taste of MSG solutions was greatly increased if ribonucleotides such as
GMP and IMP were mixed with MSG. Synergism between MSG and ribonucleo-
tides was extensively investigated by Yamaguchi (1967). She demonstrated that the
relationship between the proportion of IMP in a mixture of MSG+IMP and its per-
ceived intensity was bell-shaped (Fig. 2.7). The synergistic effect between MSG
and IMP can be expressed by the following formula:
y u v,
Fig. 2.7 Relationship between umami intensity and proportion of inosine 5′-monophosphate
(IMP) in a mixture of monosodium L-glutamate (MSG) and IMP. (Image from Umami Information
Center website, https://www.umamiinfo.jp/what/attraction/discovery/, modified from
Yamaguchi (1967))
where u and v are the concentrations (g/dl) of MSG (u) and IMP (v) in the mixture,
γ is a constant (1218), and y is the equi-umami concentration of MSG alone.
In humans, umami synergism may contribute to sensitivity to ribonucleotides,
because human saliva contains a small amount of glutamate. To test this hypothesis,
the detection threshold of IMP was investigated in the presence of MSG at various
concentrations, and it was estimated that 0.63 ppm MSG, which is lower than sali-
vary glutamate, was required to affect the detection threshold of the IMP anion
(Yamaguchi, 1991). Thus, salivary glutamate might affect sensitivity to ribonucleo-
tides, which may be based on the synergism between these substances. Synergism
between MSG and ribonucleotides has been observed in various animal species (see
Sect. 2.3). More recently, a molecular mechanism for umami synergism has been
elucidated: the TAS1R1 + TAS1R3 umami receptor is the site responsible for syn-
ergism (Zhang et al., 2008; see Chap. 1).
2.4.3 Long-Lasting
One of the unique characteristics of umami taste is that it is long-lasting, which may
be characterized as “persistency” or “aftertaste.” Time-dependent perception of
taste intensity was investigated in healthy subjects (Yamaguchi, 1998), who were
asked to keep a taste solution in their mouth for 20 s and then expectorate it. Taste
intensity was evaluated up to 100 s thereafter. When subjects sipped and
2 Umami and MSG 31
10
9
Sample
8
7
Taste intensity
6
5
4
3
2
1
Expectorate
0
0 120 240 360
Time (sec)
MSG(30mM) NaCl(132mM)
IMP(20mM) Tartaric acid(1.88mM)
Fig. 2.8 Successive time-intensity curves in response to the umami taste of monosodium
L-glutamate (MSG) and inosine 5′-monophosphate (IMP), the salty taste of NaCl, and the sour
taste of tartaric acid. (Modified from Yamaguchi (1998))
expectorated salty (NaCl) or sour (tartaric acid) solutions, the taste intensity of these
solutions rapidly decreased (Fig. 2.8). In contrast, a decrease in the taste intensity of
umami solutions (MSG and IMP) after expectorating was considerably slower. This
long-lasting effect of umami taste is concentration dependent: when the concentra-
tion of umami substances was increased, the duration of aftertaste became longer
(Kawasaki et al., 2016). This long-lasting aftertaste may explain why umami taste
has been described as persistent. Such long-lasting effects of umami taste may be
explained in part by umami signals from the larynx and the pharynx region, that is,
“stimulation in the throat.”
In mice, whole-nerve recordings from the superior laryngeal nerve innervating
the larynx demonstrated that the superior laryngeal nerve showed large responses to
MSG in a concentration-dependent manner (Arai et al., 2010). Because NaCl stimu-
lation caused a concentration-dependent decrease in responses of the superior laryn-
geal nerve, responses to MSG must be elicited by the glutamate ion, not the sodium
ion. The pharynx is innervated by the pharynx branch of the glossopharyngeal
nerve. Whole-nerve recordings from the pharynx branch of the glossopharyngeal
nerve in mice showed that umami substances such as MSG, IMP, and MSG+IMP
elicited greater responses than water stimulation, which also induced large responses
of the pharynx branch of the glossopharyngeal nerve (Kitagawa et al., 2007). In the
same manner as for the superior laryngeal nerve, NaCl stimulation elicited weaker
responses in the pharynx branch of the glossopharyngeal nerve. Therefore, responses
to MSG, IMP, and MSG+IMP were likely elicited by the glutamate and/or inosinate
ion, not the sodium ion. Interestingly, responses to MSG+IMP were almost the
same as the sum of responses to MSG and IMP, suggesting that there is no umami
synergism in the pharynx region. Thus, detection mechanisms for umami com-
pounds may be different in the oral cavity than in the pharynx.
Imamura and Matsushima (2013) identified substances in soy sauce that sup-
press this umami aftertaste. They found that polysaccharides with molecular weight
between 44,900 and 49,700 suppressed umami aftertaste. Although the mechanism
for this suppression was not elucidated, these data may indicate the existence of
receptor(s) for umami aftertaste other than TAS1R1 + TAS1R3. In summary, umami
signals from the larynx and pharynx region may contribute to the aftertaste of
umami, but this possibility should be verified with further studies.
2.4.4 Saliva Secretion
Oral taste stimulation induces saliva secretion. The volume of saliva secretion dif-
fers according to taste quality. Similar to other tastants, umami substances also
induce saliva secretion. Horio and Kawamura (1989) examined saliva secretion
from the parotid gland in response to taste stimuli in humans. Among the taste
stimuli used, tartaric acid (0.01 M) induced the largest saliva secretion; saliva secre-
tion by umami tastants such as MSG (0.1 M), IMP (0.1 M), and GMP (0.1 M) was
similar to that induced by other tastants, including NaCl (0.1 M), sucrose (1 M), and
quinine (0.0005 M). They also examined regional differences between the anterior
and posterior part of the tongue and reported that umami stimulation of the posterior
part of the tongue tended to be more effective than that of the anterior tongue,
although there was no statistically significant difference.
Other researchers have investigated saliva secretion by umami stimuli. Hodson
and Linden (2006) examined parotid saliva flow induced by taste stimuli in humans.
They demonstrated that the parotid saliva flow induced by MSG showed a dose-
dependent response and that the overall order of relative saliva flow induced by taste
stimuli was sour (citric acid) > umami (MSG) > salty (NaCl) > sweet (sucrose) ≥ bit-
ter (magnesium sulfate). Sato-Kuriwada et al. (2018) demonstrated a similar result
by examining taste-induced saliva secretion from the labial minor salivary gland;
umami and sour tastes evoked greater saliva secretion than did the other tastes. They
also showed greater saliva secretion by MSG+IMP than by MSG or IMP alone.
These studies suggest that oral umami stimulation causes greater saliva secretion
than do sweet, salty, and bitter stimulation.
Saliva secretion induced by umami stimuli may correlate with umami sensitivity
in humans. Pushpass et al. (2019) investigated the effect of older age on subjective
(perception) and objective (stimulated saliva response) measures of stimulants for
transient receptor potential channels (capsaicin, menthol), odors (menthol odor),
and basic tastants (caffeine, MSG). In this study, both perceived intensity of umami
stimulation and saliva secretion induced by umami stimulation were lower in older
subjects (>60 years) than in young subjects (18–30 years). These data indicate that
higher umami sensitivity may lead to greater saliva secretion by umami stimuli.
2 Umami and MSG 33
However, other reasons associated with human aging may underlie these results. In
addition, saliva secretion induced by umami stimuli may be long-lasting just as
umami taste perception is. Uneyama et al. (2009) demonstrated the time course of
saliva secretion after taste stimulation in healthy adult subjects. In the case of sour
stimulation (3.8 mM citric acid), saliva secretion returned to the basal level about
3 min after taste stimulation. In contrast, saliva secretion induced by umami stimu-
lation (100 mM MSG) was long-lasting, continuing for more than 10 min. Therefore,
the total amount of saliva secretion within 10 min after umami stimulation was
significantly greater than that after sour stimulation. Such an effect of umami taste
on salivation might be helpful in maintaining the oral mucosal integrity in patients
with dry mouth.
2.4.5 Mouthfullness
As described earlier in this chapter, characteristic descriptions of umami taste often

include such words as mouthfullness and persistency. These same words are elicited
by the addition of some flavor compounds named kokumi, a Japanese word literally
meaning “rich taste.” Kokumi is characterized by thickness, continuity, and mouth-
fullness in the flavors and textures (Ueda et al., 1990). By adding a water extract of
garlic to umami solutions, kokumi flavors were clearly recognized by panelists
(Ueda et al., 1990). By chromatographic separation of garlic extracts, the key com-
pounds were determined to be sulfur-containing components, such as alliin.
Many compounds are thought to impart kokumi flavor. One of the recognized
compounds found in foods that elicit kokumi flavor is glutathione (Ueda et al.,
1997). Yamamoto et al. (2009) tested the effect of glutathione on taste responses in
mice. In short-term and long-term behavioral experiments, mice showed greater
preference to IMP or MPG + IMP when glutathione was added to these solutions.
In a conditioned taste aversion paradigm, mice conditioned to avoid MPG general-
ized this response moderately to glutathione, whereas glutathione aversion did not
generalize to MPG. Gustatory nerve recordings showed synergism between IMP
and glutathione but not between MPG and glutathione. Thus, glutathione increased
preference for umami solutions containing IMP in mice. In humans, the taste inten-
sity of MSG+IMP+NaCl solution was significantly increased by the addition of
glutathione. Kokumi qualities (thickness, continuity, and mouthfullness) were also
increased by addition of glutathione added to salty, sweet, or umami solutions (Ueda
et al., 1997; Ohsu et al., 2010). Furthermore, sensory identification of MSG+NaCl
as meaty and long-lasting was increased by addition of glutathione, and an increase
in central nervous system activation attributed to MSG+NaCl+glutathione com-
pared with MSG+NaCl alone was observed in the left ventral insula in functional
MRI experiments (Goto et al., 2016). These data indicate an interaction between
umami (also sweet and salty) and kokumi.
The receptor for kokumi is believed to be the calcium-sensing receptor CaSR,
since agonist activities for CaSR correlated well with kokumi intensity (Ohsu et al.,
2010). CaSR was found in a subset of taste cells that did not express the umami and
sweet taste receptor component TAS1R3, and these cells were activated by agonists
for CaSR, including glutathione (San Gabriel et al., 2009; Maruyama et al., 2012).
Thus, umami and kokumi appear to be detected by a different subset of taste recep-
tor cells. The interaction site for umami and kokumi still has not been elucidated, but
further studies should reveal the mechanisms for such interactions.
2.4.6 Satisfaction
Another description often used for MSG flavor is “satisfaction.” This feeling not
only may depend on oral sensation but may also include information from the
throat and the gastrointestinal tract. As mentioned above, some neural information
for glutamate arises from the pharynx and larynx region, which contains taste buds
(taste cells). Furthermore, a characteristic type of cell called the solitary chemo-
sensory cell (SCC) exists in the throat (and nasal epithelium and trachea). These
cells can detect chemical substances in a manner similar to taste receptor cells
(Tizzano et al., 2011). They express the umami taste receptor TAS1R1 + TAS1R3,
although the chemosensitivity of the umami receptor in SCCs has not been eluci-
dated. It was reported that activation of SCCs leads to the release of acetylcholine,
which stimulates trigeminal nerve fibers that innervate the SCCs (Saunders et al.,
2014). Therefore, glutamate may interact with somatosensory fibers, which may
contribute to the sensations of “persistence” and “satisfaction.” After ingestion,
glutamate could enter the gut and activate umami receptors in the gastrointesti-
nal tract.
Supporting this idea, MSG infusion into the mouth, stomach, and duodenum of
rats increased afferent activity in the vagal gastric and celiac nerves (Niijima,
2000), suggesting transmission of neural information about MSG from the stom-
ach and the gut. In the gut, the umami receptor component TAS1R3 was reported
to be expressed in ghrelin-positive endocrine cells (Vancleef et al., 2018). The
ghrelin receptor is reported to be expressed in dopaminergic neurons in the ventral
tegmental area, which is involved in brain reward circuits (Zigman et al., 2006).
Therefore, activation of such reward systems in the brain by ghrelin could contrib-
ute to the sensation of “satisfaction” induced by glutamate intake. Further, gluta-
mate may stimulate umami receptors in the intestine. The umami receptor
TAS1R1 + TAS1R3 and cholecystokinin (CCK) are coexpressed in the same endo-
crine cells of mouse proximal intestine (Daly et al., 2012). They also found that
stimulation of L-amino acids, including glutamate, induced CCK release from an
STC-1 enteroendocrine cell line. CCK acts as a satiety hormone, suppressing food
intake. Thus, CCK-mediated humoral and neural signals induced by glutamate
stimulation in the intestine could also be involved in the sensation of “satisfaction”
induced by glutamate ingestion.
2 Umami and MSG 35
2.5 Conclusion and Perspective
The first paper on umami, published by Kikunae Ikeda over 100 years ago, described
many basic properties of umami taste. Subsequent studies conducted by many
researchers around the world supported and expanded Ikeda’s original observations.
However, the establishment of the scientific concept of umami taste was not achieved
until the first international symposium on umami in Hawaii in 1985. Now, umami
taste is recognized worldwide, and studies on MSG and umami taste continue to
increase. But there are still many questions on umami taste that we need to tackle,
some of which were also raised at the 100th anniversary symposium of umami dis-
covery (Beauchamp, 2009). Many of these questions, and avenues to pursue them,
are discussed in the following chapters in this volume.
Acknowledgments We thank Dr. Gary K. Beauchamp and Dr. Ana San Gabriel for helpful com-
ments, suggestions, and editing of this chapter.
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Chapter 3
Umami Taste Signaling from the Taste Bud
to Cortex
Eugene R. Delay and Stephen D. Roper
Umami is the meaty or savory taste evoked by certain amino acids present in foods,
especially monosodium glutamate (MSG) (Fig. 3.1). It is now recognized as one of
five (and possibly more) basic taste qualities that influence nutritional intake in a
wide range of animals, including humans (Roper & Chaudhari, 2017). Umami taste
is thought to signal the presence of dietary protein. In small quantities, MSG
enhances flavor and increases the palatability of food and thus food intake. This
effect gives umami a potentially important role in regulating nutritional balance
and, consequently, in maintaining health (Bellisle, 1998, 1999). As a taste quality,
umami has been recognized for over a century by Eastern cultures but only recently
has been studied by Western society. As discussed below, MSG in Western society
has had a checkered history as a taste stimulus and still is viewed by many as an
unacceptable food additive. This history, however, triggered research that has
advanced our understanding of the gustatory system, for example, identifying the
first mammalian taste receptor (Chaudhari et al., 1996, 2000) and providing the
basis for identifying cortical structures and their functions underlying cognitive sys-
tems that regulate food-directed behavior. Consequently, this chapter provides an
overview of our understanding of the mechanisms by which umami taste stimuli are
detected and subsequent signals are processed. This knowledge can be of funda-
mental importance to healthcare professions as well as to basic sciences.
E. R. Delay (*)
Regis University, Denver, CO, USA
Emeritus Faculty at the University of Vermont in Burlington, Burlington, VT, USA
S. D. Roper
Department of Physiology and Biophysics and the Department of Otolaryngology, Miller
School of Medicine, University of Miami, Coral Gables, FL, USA

https://doi.org/10.1007/978-3-031-32692-9_3
44 E. R. Delay and S. D. Roper
Fig. 3.1 Three different views of monosodium glutamate (MSG), showing a sodium ion (blue)
near the bonding carboxylic acid. (Spacing-filling models from BioTopics. http://www.biotopics.
co.uk/JmolApplet/gludisplayhalos.html)
3.1 A Brief History of Umami
Until the opening of the twentieth century, the history of umami was in the realm of
culinary arts. The name umami itself did not yet exist. Kitchen lore included the use
of seasonings, broths, mushrooms, meat and fish extracts, and other savory ingredi-
ents to enhance the palatability of prepared foods. Kikunae Ikeda, a distinguished
food chemist working at Imperial University of Tokyo’s College of Science at that
time, conjectured that a fundamental taste quality was the basis for the savory or
meaty taste sensation in fish, meat, and most notably, broth prepared from bonito or
dried seaweed. Ikeda reasoned that this taste quality was distinct from the traditional
four basic tastes: sweet, sour, salty, and bitter. He prepared aqueous extracts of dried
seaweed, which he selected as the primary material because its protein content
could readily be removed. After an extensive series of extractions to remove the
unexpectedly high amounts of mannitol (200 g from 1 kg of dried seaweed) and the
anticipated sodium chloride and potassium chloride from the seaweed extract, Ikeda
was able to crystallize a miniscule amount of the amino acid glutamic acid. When
he tasted a sample, it evoked a weak sourness along with a strong savory taste he
named umami (Ikeda, 1909, 2002).
Glutamic acid (glutamate) was not unknown to food chemists at the time. This
amino acid had first been isolated from wheat gluten (hence the name) in 1866 by
H. Ritthausen. Previewing its later discovery by K. Ikeda as the prototypic umami
stimulus, Ritthausen noted that glutamate elicits a unique “meaty” aftertaste:
“…Entfernt an den Nachgeschmack einer geringen Menge von concentrirtem
Fleischextract” (“[glutamate has]…somewhat of an aftertaste of a small amount of
concentrated meat extract”; Ritthausen, 1866). Other chemists cataloged glutamic
acid as having an unpalatable, weakly sour taste (Fischer, 1906). However, Ikeda
recognized that the (neutral) sodium, potassium, and calcium salts of glutamic
acid have an intense umami taste. In his remarkable 1909 paper, where he outlines
his discovery of the savory taste umami, Ikeda commented that the preference for
glutamate conceivably evolved with the consumption of meat, which always con-
tains varying amounts of glutamate (Ikeda, 1909, 2002). He compared this with an
evolution of sweet taste, which is imparted by sugars in nutritious vegetables
and fruits.
3 Umami Taste Signaling from the Taste Bud to Cortex 45
According to the historian Jordan Sand (2005), Ikeda, who had been trained in
Germany, was strongly influenced by Justus von Liebig, a leader in food chemistry.
In 1840 von Liebig had extracted the essence of meat and invented a beef extract
that later became the world-famous product Oxo. Sand mentions that von Liebig’s
beef extract “fed German armies” in the nineteenth century, and Oxo was widely
used in military rations in World War I. Half a century later, after Ikeda had discov-
ered, patented, and promoted MSG, this additive was also incorporated into military
rations for the Japanese army and, after World War II, for US Armed Forces MREs
(meals ready to eat). In 1952 MSG was included in the Marine Corps Recipe Manual
(US Marine Corps, 1952) which, for example, listed the additive in a recipe for
creamed beef (Fig. 3.2). Perhaps the use and acceptance of MSG in Western cuisine
after World War II was influenced by young recruits consuming this flavor enhancer
in their meals, though there is no hard evidence for this speculation (see, e.g.,
Geiling, 2013).
Ikeda was aware of the impact of his discovery of umami and its potential as a
food seasoning, like von Liebig’s meat extract. Indeed, he patented MSG in Japan,
the United States, England, and France and began producing the substance as a
seasoning named Ajinomoto (meaning “essence of aji” or “taste”). Ajinomoto,
a.k.a. MSG, was fairly quickly accepted in Japan, which was undergoing a rapid
cultural evolution in the early twentieth century. Initially, due to its cost, only the
Fig. 3.2 Recipe for creamed beef from the US Marine Corps Recipe Manual (US Marine Corps,
1952), documenting use of MSG in US Armed Forces mess halls
wealthier segments of society could afford it. However, as the twentieth century
progressed, women adopted a new domesticity geared toward scientific inventions
and discoveries and were ready to accept Ajinomoto. With continued efforts to
make MSG more affordable to the general population, in 1931 Ajinomoto was sold
for the first time in containers equivalent to saltshakers that could be placed on a
table (Sand, 2005).
Use of MSG was slower to reach the US market. However, ongoing scientific
research in an entirely different area, neuroscience, was soon to make a discovery
that put an indelible and, as has later been shown, an undeserved stigma on
MSG. Researchers studying the effects of a number of agents on hereditary retinal
dystrophy in rodents noticed that injecting monosodium glutamate into pregnant
female mice resulted in damage to the inner retina (Lucas & Newhouse, 1957), with
effects much more pronounced in the newborn mice than in the mothers. Further,
parental injections of glutamine, quinine, epinephrine, methanol, and other com-
pounds showed no similar retinal pathology. Because of the similarity between reti-
nal cells and neurons in the central nervous system (CNS), these experiments were
soon repeated by other laboratories focused on damage in the brain. A firestorm of
claims and counterclaims ensued regarding whether and how MSG injections dam-
aged brain tissue. Then in 1968, a letter appearing in the prestigious New England
Journal of Medicine claimed that MSG, a seasoning commonly used in Chinese
restaurants, seemed to cause “…numbness at the back of the neck, gradually radiat-
ing to both arms and the back, general weakness and palpitation” (Kwok, 1968).
This became known as the “Chinese restaurant syndrome.” Soon after, a pair of
letters based on anecdotal experiences—one written by a group of second-year
medical students—again in the New England Journal of Medicine stated that MSG
was at the root of Chinese restaurant syndrome (Ambos et al., 1968; Schaumburg &
Byck, 1968).1 The students even proffered therapies: Atarax (hydroxyzine, an anti-
histamine), Librium (chlordiazepoxide, an anxiolytic), and atropine (a cholinergic
receptor antagonist). A subsequent study by one of these groups published in the
leading journal Science cited experimental evidence for MSG as the cause of
Chinese restaurant syndrome (Schaumburg et al., 1969). This cemented the fate of
MSG in the eyes of many, who were convinced that the seasoning was at the root of
their bad experiences with oriental food. Subsequently, several organizations have
conducted thorough and exhaustive investigations of the dangers of ingesting
MSG. In each case, MSG was declared safe for consumption (e.g., Bellisle, 1999;
US Food & Drug Administration, 2020). Moreover, the authenticity of the initial
claim in the New England Journal of Medicine about MSG and the Chinese restau-
rant syndrome has been disputed (Blanding, 2019; Glass, 2019). Despite all this,
and against overwhelming evidence for its safety, the questionable reputation of
MSG lingers on in the eyes of many consumers.
1
Interestingly, regarding Chinese restaurant syndrome, the lead author of one of these publications
had only a few weeks previously (and in the same journal) explicitly stated, “I don’t think the cause
is soy sauce or monosodium glutamate” (Schaumburg, 1968).
What is indisputable, however, is that the sodium salt of L-glutamate (MSG) is

found naturally in abundance in many common foods, such as cheeses (especially
Parmesan cheese), meats, fish, and vegetables (such as tomatoes, mushrooms, egg-
plant). Also, it is unassailable that MSG, especially in combination with other foods,
is a preferred taste for humans and other animals. Thus, the history of umami is not
one of the discoveries of a new taste but the story of how an existing taste has been
identified, popularized, scrutinized, and criticized.
3.2 Umami Psychophysics: Humans and Rodents
Research has given us some understanding of the psychophysical properties of

umami taste that gives it the ability to influence ingestive behavior and nutritional
regulation. Among the most fundamental properties of a sensory system is its sensi-
tivity to stimulus intensity. In taste, detection thresholds establish the minimum
intensity (concentration) at which the presence of a substance can be sensed.
Knowledge of detection thresholds provides important standards for diagnosing
chemosensory disorders and studying physiological and molecular mechanisms.
In general, detection thresholds for glutamate appear to average between 0.5 and
2 mM in human adults (Yamaguchi & Kimizula, 1979; Schiffman et al., 1981),
which is unaffected by the concentration of sodium (Na+) (Yamaguchi, 1991).
Monopotassium glutamate and monosodium aspartate, which are also umami com-
pounds (Maga, 1983), have similar detection thresholds (Schiffman et al., 1981;
Yamaguchi, 1991).
Inosine 5′-monophosphate (IMP) and 5′-guanosine monophosphate (GMP),
which are catabolic products of nucleic acids that are often found alongside gluta-
mate in many meats and vegetables, are also flavor enhancers that elicit an umami
sensation. The detection threshold for IMP (a disodium salt) is in the same range as
that for MSG, but unlike MSG, its value is affected by the presence of Na+
(Yamaguchi, 1991). Mixtures of MSG plus IMP or GMP are synergistic, that is, are
capable of reciprocal increases in sensitivity. Indeed, one of the defining properties
of umami taste is a synergy between MSG and IMP or GMP (Yamaguchi &
Kimizula, 1979). Either nucleotide can intensify the umami sensation of MSG and
other amino acids in a nonlinear manner (Rifkin & Bartoshuk, 1980; Yamaguchi,
1991; Kawai et al., 2002). Subthreshold concentrations of IMP lower the detection
threshold for MSG taste by nearly 100-fold, and conversely, the threshold of IMP is
lowered by MSG (Yamaguchi, 1991).
Another important property of sensory systems is recognition threshold, the min-
imum intensity at which a stimulus can be identified, not merely detected, and
begins to exert motivating influences over behavior (Halpern, 1997). Recognition
thresholds typically are higher than detection thresholds. Yamaguchi (1991) found
that about 50% of subjects were able to identify the umami taste of MSG at a con-
centration twice its detection threshold. In contrast, identifying the umami taste of
IMP required a concentration four times its detection threshold. In comparison,
recognizing the salty taste of NaCl required concentrations more than ten times its
detection threshold. As might be expected, mixing MSG and IMP significantly low-
ers the recognition threshold for umami (Shigemura et al., 2009). It may be impor-
tant to note that the concentration of glutamate and IMP in natural products varies
widely, from below to well above recognition thresholds (Giacometti, 1979;
Ninomiya, 2003). Moreover, either compound might influence taste perception by
interacting with yet other food substances at or near recognition thresholds. An
important consideration is that genetic variations in umami taste receptors appear to
directly affect sensitivity and recognition thresholds for umami compounds in
humans and mice (Raliou et al., 2009; Shigemura et al., 2009).
Evidence from a variety of sources supports Ikeda’s initial observation that glu-
tamate elicits a unique taste quality (Ikeda, 1909, 2002). For example, human sub-
jects use different verbal qualifiers to describe glutamate taste compared to the other
four basic tastes (Yamaguchi, 1991; Hettinger et al., 1996). This effect crosses cul-
tural boundaries (Yamaguchi, 1991; Prescott, 1998). Interestingly, MSG and other
umami compounds at concentrations found in food additives are hedonically posi-
tive and are typically described as “savory” or “meaty,” but high concentrations of
MSG alone are not preferred by humans (Schiffman et al., 1981; Okiyama &
Beauchamp, 1998).
The perceived taste of glutamate salts is often complex due to the presence of
sodium or other cations. The most important attribute of glutamate and other umami
compounds in fact may be their ability to enhance the palatability of other food
components. When added to solutions containing compounds that elicit a single
basic taste (e.g., sucrose/sweet or quinine/bitter), MSG has little effect on the qual-
ity or intensity of the taste (Yamaguchi & Kimizula, 1979). However, when MSG is
added to soup broth, potatoes, or other food items, subjects find them much more
palatable and exhibit eating behaviors consistent with an increase in hedonic value
(e.g., increase eating rates or shorter between-bite pauses), especially if paired with
a novel flavor or with the odor of a savory vegetable (Bellisle & Le Magnen, 1981;
Rogers & Blundell, 1990; Okiyama & Beauchamp, 1998; Prescott & Young, 2002;
Prescott, 2004; McCabe & Rolls, 2007).
Studying perceptual experiences of nonhuman animals is a challenging but
important endeavor for chemoreception sciences. Much of our understanding of
cellular and molecular mechanisms of taste transduction, including umami taste, is
based on research with nonhuman species. Direct comparisons with human percep-
tual experiences are difficult at best, but a number of methods have been used to
develop psychophysical profiles of gustatory phenomena (e.g., Spector, 2003).
Comparing taste profiles from animal studies with human taste profiles for the same
umami substances reveals striking similarities and some important species-specific
characteristics. Taste sensitivity of rodents for glutamate and L-aspartate is compa-
rable to that of humans. For example, detection thresholds are between 1 and 4 mM
for rats (Stapleton et al., 2002; Taylor-Burds et al., 2004) and between 0.01 and
2.5 mM for mice2 (Stapleton et al., 2002; Mukherjee & Delay, 2011). Recognition
threshold in rats for glutamate taste is between 5 and 10 mM (Yamamoto et al.,
1991; Chaudhari et al., 1996; Stapleton et al., 1999; Heyer et al., 2003), whereas
mice have slightly higher thresholds. Interestingly, at low, near-threshold concentra-
tions of MSG, rodents can confuse the taste of glutamate with sucrose (Yamamoto
et al., 1991; Chaudhari et al., 1996; Stapleton et al., 1999; Heyer et al., 2003). Rats
and mice generally show a natural preference (positive valence) for MSG, IMP, and
other L-amino acids, even at concentrations that humans find unpleasant (Pritchard
& Scott, 1982; Iwasaki et al., 1985; Delay et al., 2000; Ruiz et al., 2003; Wifall
et al., 2007), although some of this may be related to postingestive effects (Ackroff
& Sclafani, 2016). The perceptual uniqueness of MSG has been demonstrated in
mice, which did not generalize a conditioned taste aversion between MSG and the
other four basic tastes (Ninomiya & Funakoshi, 1987). Synergy between MSG and
IMP in rats has also been reported with brief-access taste tests (Yamamoto et al.,
1991; Delay et al., 2000). In addition, it should be noted that detection thresholds
for a number of umami stimuli can be influenced by a variety of factors such as
temperature, pH, age, diet, and other variables that also are important in food prepa-
ration and perception (Barragan et al., 2018; Green et al., 2016; Jeon et al., 2021;
Ma et al., 2020; Zhong et al., 2015.
3.3 Overview of Tongue and Gustatory System
Taste sensations are generated in the oral cavity, primarily from taste buds on the
tongue, and are transmitted to higher regions in the brain for analysis and interpre-
tation. An overview of the gustatory system is shown in Fig. 3.3. Briefly, taste
stimuli are transduced into afferent signals by specialized sensory cells in taste
buds embedded in the oral epithelium. These gustatory sensory cells transmit
these signals to primary sensory afferent fibers of cranial nerves 7, 9, and 10 (CN7,
CN9, CN10, respectively) that project into the hindbrain to the nucleus of the soli-
tary tract (NST). In rodents, afferent signals are then sent to the parabrachial
nucleus (PBN) in the pontine area and from there to the ventroposterior medial
parvicellular (VPMpc) nucleus of the thalamus or to subcortical structures in the
lateral hypothalamus, amygdala, and other structures. Thalamic signals are then
transmitted to the insular cortex and other cortical areas. In primates, neurons in
the NST project to the thalamus and thalamic neurons project to neurons in the
primary gustatory cortex, the frontal operculum/insula (FOI), where the ability to
perceive tastes (e.g., sweet and bitter) is thought to occur. Subsequent processing
2
Specifically, C57BL/J6 mice, a mouse strain often used for genetic manipulations such as gene
deletions (knockout) or labeling of proteins involved in taste transduction. Different strains of mice
have notoriously different taste thresholds (Bachmanov et al., 2016). Other researchers have
reported different threshold estimates (Nakashima et al., 2012; Blonde et al., 2018; Smith &
Spector, 2014).
Fig. 3.3 Overview of rodent and human gustatory systems. (a and b) Schematic diagram of the
major structures of the gustatory system (a), including sensory input from taste cells and taste buds
and from the gut to the nucleus of the solitary tract (NST) and the principle ascending pathways to
cortical and subcortical structures that influence taste-directed behavior. In the rodent (b), input
from the tongue via cranial nerves 7, 9 and 10 (CN7, CN9, CN10) goes to the NST, whose output
goes to the parabrachial nucleus (PBN) and then to the ventroposterior medial (parvicellular)
(VPMpc) nucleus of the thalamus. Other subcortical structures, such as the lateral hypothalamus
(LH) and amygdala (Amy), also receive taste information from the PBN. From the thalamus, taste
information is then sent to the insular region of the cortex and to other cortical areas, such as the
cingulate gyrus (CG). BG, basal ganglia; FOI, frontal operculum/insular area; OFC, orbitofrontal
cortex. (c–e) Flow of taste signals from the cranial nerves to areas of the cortex that process and
subsequently influence taste-directed behavior in humans. (c) A view of the medial aspects of the
taste system showing the ascending flow of afferent signals through the NST to the VPMpc. (d)
From a dorsal perspective of the right hemisphere, taste signals go from the VPMpc to the FOI, the
primary taste cortex. These signals are then sent to the OFC and other cortical areas, such as the
CG, and subcortical structures, such LH, Amy, or BG. (e) A lateral view of the right hemisphere
with the approximate locations of the FOI and the OFC identified
by association cortices such as the orbitofrontal cortex (OFC) and cingulate gyrus
(CG) contributes to higher-order cognitive processes involved in taste-directed
behavior. In humans, it appears that umami taste signaling from the tongue to the
cortex is predominantly ipsilateral (Iannilli et al., 2012). Below a fuller descrip-
tion of each step in the taste pathway is presented.
3.4 Receptors
The concept that there are specific cell-membrane binding sites for sweet—a
glucophore-binding site (Shallenberger & Acree, 1967)—and for salt, a sodium
receptor (Beidler, 1954), dominated ideas about a molecular basis of taste reception
in the middle of the twentieth century. These ideas were generalized to other quali-
ties such as the “acidophore” (a hydrated proton) receptor for sour (Shallenberger,
1993). Yet these concepts remained theoretical, and the actual identity of membrane
surface molecules responsible for interacting with taste compounds was elusive.
Only sometime later did researchers begin in earnest to study the molecular basis of
umami taste, because of the lack of acceptance of umami as a separate, basic taste.
Annick Faurion was an early pioneer in the efforts to identify umami receptors. She
surmised that umami taste receptors may be akin to the newly characterized NMDA
glutamate synaptic receptors found in the brain (Faurion, 1991). Initial efforts to test
this experimentally suggested that there were indeed NMDA-like receptors in mem-
branes isolated from fish lingual tissues rich in taste buds (Brand et al., 1991; Teeter
et al., 1992), but pinpointing the results specifically to taste cells was not possible in
those experiments.
A major breakthrough occurred when metabotropic synaptic glutamate receptors
(G-protein-coupled receptors, GPCRs) were cloned and identified in the brain
(Houamed et al., 1991; Masu et al., 1991). There was reason to believe that taste
transduction might involve GPCRs because of the early efforts of Naim et al. (1991)
showing that in taste tissues, sweet taste generated cAMP, a key second messenger
for many GPCRs. Additionally, a taste-specific Gα protein had been cloned and
characterized (McLaughlin et al., 1992), reinforcing the notion that taste involved
GPCRs. By analogy, it was believed that umami taste might also involve GPCRs.
Chaudhari and colleagues identified a novel, truncated metabotropic synaptic gluta-
mate receptor, taste-mGluR4, in rat taste buds and postulated that this molecule
might serve as an umami receptor (Chaudhari et al., 1996, 2000). Taste-mGluR4 fit
all the requirements for a candidate taste receptor: (a) it was present selectively in a
small subset of taste bud cells (Chaudhari & Roper, 1998; Yang et al., 1999), and (b)
when expressed in a heterologous cell line (CHO cells), the receptor conferred glu-
tamate sensitivity at taste-appropriate concentrations (Chaudhari et al., 2000).
mGluR4 knockout (KO) mice (mutant mice lacking a functional mGluR4 gene)
showed abnormal glutamate taste behavior, but the results were enigmatic: they had
reduced taste nerve responses to MSG compared to wild-type mice (Yasumatsu
et al., 2015) but showed increased, not decreased, preference for umami taste solu-
tions (Chaudhari & Roper, 1998). This taste behavior in mGluR4 KO mice could
perhaps be interpreted as due to a decline or muting in umami taste sensations, driv-
ing the mutant mice to consume more of the solution to obtain reinforcement. Yet,
interpreting the effects of a global knockout (mGluR4 KO mice) is complicated by
the fact that this receptor has widespread functions in neural circuitry in the brain;
its deletion likely affects many cognitive processes, not merely gustation.
Soon after the discovery of taste-mGluR4, other taste-specific umami receptors
were cloned and identified in mouse taste buds. These receptors were also GPCRs
and consisted of two different gene products, T1R1 and T1R3, combined into a
heterodimer (Nelson et al., 2002; Zhao et al., 2003). The T1R1 + T1R3 heterodimer
had similar properties to taste-mGluR4: the molecules were found in a subset of
gustatory receptor cells, and expression in heterologous cells conferred sensitivity
to glutamate and other amino acids. Importantly, Zhao et al. (2003) reported that
mice lacking T1R1, T1R3, or the T1R1 + T1R3 receptor heterodimer were taste
blind to MSG. However, these findings have been challenged. T1R3 KO mice had
only slightly elevated MSG detection thresholds (Damak et al., 2003; Delay et al.,
2006), challenging the notion that T1R1 + T1R3 receptor heterodimers are the only
umami taste receptors and supporting important roles for mGluR4 and other gluta-
mate receptors in umami taste (Yasuo et al., 2008; Delay et al., 2009; Yasumatsu
et al., 2009; Kusuhara et al., 2013; Blonde & Spector, 2017; Blonde et al., 2018).
More recently, another candidate umami receptor, a truncated form of the
metabotropic glutamate 1 receptor, taste-mGluR1, has emerged (San Gabriel et al.,
2005; San Gabriel et al., 2009). Mutant mice lacking mGluR1 have not yet been
tested for taste behavior, but the interpretation of these data would be subject to the
same reservations as for taste behavior assays in mGluR4 KO mice: mGluR1 is an
important synaptic receptor in the brain, and behavioral alterations might be wide-
spread in mGluR1 KO mice, as was described above for mGluR4 global knockout.
In summary, at least four different candidate umami taste receptors have been put
forward: NMDA-like, taste-mGluR4, taste-mGluR1, and the T1R1 + T1R3 het-
erodimer. No strong experimental evidence for NMDA-like umami receptors in
taste buds has yet been found, and the bulk of evidence favors the other three recep-
tor candidates.3 Thus, multiple receptors—T1R1 + T1R3, mGluR1, and mGluR4—
may underlie umami taste.
3.5 Structure and Function of Umami Receptors
All the receptors identified to date for umami taste transduction are class C GPCRs.
This class of GPCRs is characterized by an extensive extracellular domain, constitu-
tive dimerization, and an unusual N-terminal bilobed ligand-binding region that
resembles a Venus flytrap, hence its name: the Venus flytrap (VFT) domain. By
analogy with the sweet taste receptor heterodimer, T1R2 + T1R3 (Nelson et al.,
2001), T1R umami receptors were shown to be heterodimers of T1R1 + T1R3 (Li
et al., 2002; Nelson et al., 2002) (Fig. 3.4). Further, by analogy with synaptic
mGluRs, the glutamate binding site for T1R1 + T1R3 was shown to reside in the
VFT domain of T1R1 (Zhang et al., 2008; Lopez Cascales et al., 2010; Roura et al.,
2011; Toda et al., 2013). IMP interacts with a nearby site to stabilize the closed and
active VFT domain occupied by glutamate (Zhang et al., 2008), explaining the abil-
ity of IMP to enhance umami taste. Compounds that bind to a transmembrane region
of T1R3 also modify umami taste. Examples include lactisole, a sweet taste inhibi-
tor that interferes with umami taste (Xu et al., 2004), and cyclamate, an artificial
sweetener that enhances umami responses (Zhang et al., 2008).
3
Ionotropic glutamate receptors, including NMDA receptors, are expressed on one of the types of
taste bud cells (specifically, Type III cells—those that respond to sour taste; Roper & Chaudhari,
2017). However, instead of participating in the initial transduction of glutamate taste, these recep-
tors appear to be involved in signal processing and feedback synaptic circuitry within taste buds
(Vandenbeuch et al., 2010; Huang et al., 2012). The presence of these synaptic glutamate receptors
may explain early reports claiming the expression of NMDA receptors in taste buds as evidence for
umami transduction via these receptors.
A Ba b c
glutamate,
IMP glu
glu
lactisole,
cyclamate
IMP
T1R1 T1R3
Fig. 3.4 T1R1 + T1R3 heterodimer umami taste receptor. (a) Glutamate and IMP bind to the large
extracellular Venus flytrap domain of T1R1 in the dimeric umami taste receptor. (Modified from
Laffitte, Neiers et al., 2014; Roper, 2020). (b) Molecular mechanism of the umami receptor: rib-
bon-band representation of the Venus flytrap motif on the T1R1 + T1R3 umami receptor in three
situations—no bound ligands (a), binding of glutamate (glu) (b), and binding of both glutamate
and GMP (c). (Modified from Mouritsen et al. (2013))
mGluR4 taste-mGluR4
Fig. 3.5 The mGluR umami taste receptor (right) is a truncated splice variant of synaptic mGluR
(left). The heavy black line shows the Venus flytrap motif, significantly truncated in taste-mGluR4.
(From Chaudhari et al. (2000))
As noted above, the metabotropic umami receptors mGluR1 and mGluR4 found
in taste buds are class C GPCRs. These umami taste receptors are distinct from their
synaptic glutamate receptor equivalents. Specifically, taste-mGluR1 and taste-
mGluR4 umami receptors are N-terminal truncated variants of synaptic mGluR1
and mGluR4 receptors (Fig. 3.5). Interestingly, the truncation eliminates about half
the VFT domain, the known glutamate binding region for synaptic mGluRs (O’Hara
et al., 1993). Structure-function analyses of glutamate binding domain(s) have not
been carried out for taste-mGluR1 or taste-mGluR4. Much less is known about
whether and how MSG activates these mGluR umami receptors. Further, although
synaptic mGluRs form dimers (Kunishima et al., 2000), it is not known whether the
mGluR1 or mGluR4 umami taste receptor does so.
3.6 Downstream Signaling
Signal transduction downstream of the T1R1 + T1R3 umami receptor follows the
canonical GPCR-inositol trisphosphate (IP3)-intracellular Ca2+ release pathway,
extensively documented in a number of excellent reviews on taste (Kinnamon,
2009; Roper & Chaudhari, 2017; Kinnamon & Finger, 2019; Roper, 2020; Gutierrez
& Simon, 2021) (Fig. 3.6). The transduction cascade is initiated by glutamate bind-
ing to T1R1 + T1R3 on taste bud umami-sensing cells (specifically, type II taste
cells, as distinct from type I glial-like taste cells and type III sour-sensing taste cells;
see Roper & Chaudhari, 2017) activating G-proteins, initiating intracellular Ca2+
release, which activates TRPM4 and TRPM5 cation channels. The depolarization
produced by cation influx through these channels triggers action potentials in the
cell, which opens large-pore CALHM 1 and 3 ion channels that allow the release of
ATP, the principal type II cell transmitter Finger et al., 2005; Ma et al., 2018).4
Early studies also implicated a role for cAMP in the umami transduction path-
way. Glutamate stimulation of taste tissue decreases cAMP (Abaffy et al., 2003),
and genetically engineered mice lacking Gα gustducin, the G-protein that couples
taste GPCRs to cAMP metabolism, have diminished taste responses to glutamate
(He et al., 2004). The rather convoluted concept that has evolved (Clapp et al., 2008)
(Fig. 3.6, gray arrows) is that cAMP inhibits key steps in the above canonical IP3
pathway and gustducin tonically activates cAMP-dependent phosphodiesterase to
maintain cytosolic cAMP at a low level (McLaughlin et al., 1994). In this way, gust-
ducin maintains both phospholipase C β2 (PLCβ2) and inositol 1,4,5-trisphosphate
receptor, type 3 (IP3R3), in a primed and ready state (Clapp et al., 2008).
Curiously, little is yet known regarding umami transduction pathways initiated
by taste-mGluR1 and taste-mGluR4. This is an area of research that remains to be
developed.
3.7 Cranial Nerve Responses to Umami
Study of the afferent pathway gives us some novel insights into the encoding pro-
cess of umami substances. Taste buds are innervated by three cranial nerves: the
facial (CN7), glossopharyngeal (CN9), and vagus (CN10) nerves. Two branches of
4
Interestingly, unlike synaptic release elsewhere in the nervous system, ATP release in type II cells
is nonvesicular and involves only depolarization-activated CaHLM1/3 channels, independent of
intracellular Ca2+ (Nomura et al., 2020). Indeed, type II taste bud cells lack voltage-gated calcium
channels (Clapp et al., 2006).
(tonic)
Gα
PDE3 ¯ [cAMP] ¯ PKA
¯
gustducin
T1R1
+T1R3
¯
Gβg
PLC ¯
β2
IP
IP 3
3R
¯
3
[C
¯
a 2+
]
TR PM
TR
PM 4
5,
N a in
a_ fl
C
, ux
me epol’
d
mb n
ran
e
ope M1,3
CaH
ns
L
ATP
release
Fig. 3.6 Representation of the canonical G-protein-coupled receptor chemosensory transduction

cascade. Open arrows symbolize the pathway for G-protein activation that leads to intracellular
Ca2+ mobilization, depolarization (depol’n), and neurotransmitter (ATP) release. Gray arrows at
the top depict the constitutive (tonically active) Gα gustducin pathway that results in downregula-
tion of protein kinase A (PKA). Tonic activation of this pathway disinhibits key elements of the
canonical pathway: phospholipase C β2 (PLCβ2) and inositol 1,4,5-trisphosphate receptor, type 3
(IP3R3). The signal(s) that maintains constitutive Gα gustducin activation is unknown, though
taste receptor stimulation is one likely contributor (Clapp et al., 2008). PDE3, phosphodiesterase
3; IP3, inositol trisphosphate; TRPM4 and TRPM5, transient receptor potential cation channel
subfamily M, members 4 and 5; CALHM1, 3, calcium homeostasis modulator 1 and 3. (Modified
from Roper (2020))
the facial nerve, the chorda tympani and the greater superficial petrosal, innervate
taste buds in the anterior portion of the oral cavity. The chorda tympani innervates
fungiform papillae on the anterior two-thirds of the tongue and some taste buds in
foliate papillae on the lateral tongue. The greater superficial petrosal innervates
taste buds in the soft palate. The glossopharyngeal nerve innervates the posterior
third of the tongue, including taste buds in the circumvallate papillae and some in
the foliate papillae. The vagus nerve innervates taste buds and solitary chemorecep-
tors in the posterior oral cavity and throat. All of these fibers synapse in the NST,
which relays information to other structures of the CNS.
Whole-nerve and single-fiber recordings from gustatory nerves have provided

abundant evidence clarifying how umami taste signals are handled by the nervous
system. Early investigators demonstrated the synergistic interaction between MSG
and a number of 5′-ribonucleotides in rat with whole-nerve (Adachi, 1964) and
single-fiber (Sato et al., 1970) recordings of the chorda tympani. More recently,
Sako et al. (2000) found that the response to MSG was similar in the greater super-
ficial petrosal nerve and the chorda tympani of rats, including synergistic respond-
ing to mixtures of MSG and IMP. In contrast, the glossopharyngeal nerve appears to
carry a smaller umami signal, with little or no evidence of MSG-IMP synergy. The
greater contribution of the chorda tympani and the greater superficial petrosal nerves
for umami signaling were verified by the finding that rats with transections of
chorda tympani and superficial petrosal nerves were unable to learn a conditioned
taste aversion to MSG mixed with IMP to the same degree of rats with transaction
of both the glossopharyngeal and either of the other nerves (Ninomiya & Funakoshi,
1987, 1989). Nonetheless, the importance of this smaller glossopharyngeal signal
should not be ignored. If the glossopharyngeal nerve is transected, mice conditioned
to avoid MSG cannot distinguish MSG from NaCl (Ninomiya & Funakoshi, 1987,
1989), suggesting that the glossopharyngeal nerve also transmits important qualita-
tive information about glutamate taste.
Early research using whole-nerve recording methods found that MSG and other
umami substances elicited strong responses in the chorda tympani, quite similar to
responses elicited by sucrose (e.g., Sato et al., 1970) or NaCl (Yamamoto et al.,
1991). At this time, the overlap in nerve responses to umami, NaCl, and sucrose
raised questions about whether umami was a basic taste or simply a combination of
sucrose and NaCl activity. This forced researchers to compare glutamate responses
to those elicited by NaCl and sucrose to identify any unique effect attributable only
to glutamate. Single-fiber recording studies and the reduction of responses by the
sodium inhibitor amiloride and by sweet taste inhibitors have helped researchers
parse the components of MSG-evoked whole-nerve responses. Single-fiber record-
ing studies have searched for fibers that respond best to MSG and other umami
substances (so-called M-best fibers), but evidence of these fibers has been slow to
accumulate because these studies have often encountered fibers that appear to carry
signals for MSG as well as for sucrose or NaCl, especially in the chorda tympani.
Moreover, the population of M-best fibers appears to be much smaller than that for
sweet (S-best fibers) or salt (N-best fibers) stimuli. Nevertheless, there is now evi-
dence of M-best fibers in several species, such as mouse, rat, pig, dog, and chimpan-
zee (Danilova et al., 1999; Hellekant et al., 1997; Kumazawa et al., 1991; Ninomiya
& Funakoshi, 1987, 1989), but not in hamsters (Yamamoto et al., 1988). M-best
fibers respond to MSG and do not respond to sucrose. They also often show synergy
between MSG (or monopotassium glutamate) and either IMP or GMP.
While evidence of M-best fibers accumulated slowly, evidence that signaling of
umami stimuli also involves nerve fibers that respond to sucrose was discovered in
early studies (Sato et al., 1970; Ninomiya & Funakoshi, 1987, 1989), which prob-
ably explains why rodents often have difficulty discriminating sucrose and gluta-
mate at lower concentrations (Yamamoto et al., 1991; Stapleton et al., 2002; Heyer
et al., 2003). Four fiber types in the chorda tympani of mice have been identified
based on their responses to sucrose and monopotassium glutamate and evidence of

synergy when IMP is mixed with glutamate (Yasumatsu et al., 2012, 2015): M-best
fibers exhibit synergy (M1 fibers) or not (M2 fibers), and sucrose-best fibers exhibit
synergy (S1 fibers) or not (S2 fibers) when stimulated with monopotassium gluta-
mate and IMP. Subsequent studies using an array of sweet inhibitors and glutamate
agonists and antagonists determined that each fiber type appears to be activated by
a specific set of taste receptors: S1 and S2 fibers are activated by T1R receptors, M1
fibers are activated by mGluR1 receptors, and M2 fibers are activated by mGluR4
receptors (Yasumatsu et al., 2012, 2015). Thus, the density of each receptor family
along the anterior-posterior dimension of the tongue appears to influence the nature
of glutamate responses within each nerve.
Lastly, in mice, recordings from geniculate ganglion neurons that innervate taste
buds on the anterior tongue and soft palate reveal a small population of sensory
neurons that respond exclusively to MSG, presumably representing the parent neu-
rons of M-best fibers (Barretto et al., 2015; Wu et al., 2015).5 However, hierarchical
clustering of the geniculate ganglion neurons showed a good deal of overlap and no
clean separation between clusters of sucrose- and MSG-responding sensory neu-
rons (Wu et al., 2015), reinforcing the similarity between sucrose and umami tastes,
at least in rodents, and the involvement of the T1R3 monomer that is common to
both sweet and umami taste receptors.
3.8 Nucleus of the Solitary Tract and Parabrachial Nucleus
Far fewer studies have examined the response of CNS neurons to MSG, and most of
these studies have been conducted in rats and mice. These have often compared
response patterns of neurons to equimolar concentrations of NaCl, sucrose, and
MSG. The most illuminating of these studies have used the salt taste inhibitor
amiloride to help dissociate the responses of Na+ and the glutamate anion. Neural
responses to umami substances from neurons located in the NST have received
some attention. In the rat, neuronal responses to 0.1 M MSG were quite similar to
responses elicited by 0.1 M NaCl (Giza & Scott, 1991; Giza et al., 1996, 1997).
However, the addition of amiloride reduced the overall response to NaCl and
changed neuronal response profiles more for NaCl than for MSG, presumably due
to a glutamate anion signal that is unaltered by the presence of amiloride.
Interestingly, profiles of neural responses in the NST of rats revealed differences in
temporal coding between sucrose and MSG taste stimuli in awake and behaving rats
(Roussin et al., 2012). This suggests that brain stem coding and transmission of
taste qualities of umami, NaCl, and sucrose may be accomplished by overlapping
populations of neurons but qualitatively distinguished by more subtle properties in
the train of action potentials.
5
Neither of these studies attempted to differentiate M1 and M2 responses. These studies used a
mixture of IMP with MSG (Wu et al., 2015) or monopotassium glutamate (Barretto et al., 2015).
The gustatory portion of the NST projects to the medial PBN in the rodent
(Norgren, 1978). In the rat, neurons in the medial PBN do not exhibit as strong a
relationship in their response to NaCl plus MSG or to sucrose plus MSG as do neu-
rons in the NST, suggesting more dissociation in the pathways carrying the afferent
signals for these stimuli (Nishijo et al., 1991). In the mouse PBN, sucrose and
umami signals appear to be processed more medially, whereas signals for other
basic tastes are processed more laterally (Tokita et al., 2012). Even so, evidence of
overlapping taste signals for sucrose, umami, and NaCl has been observed. Many of
the medially located neurons identified as sucrose-best neurons also show stronger,
synergistic responses to mixtures of monopotassium glutamate and IMP, indicating
convergence of glutamate taste signals with sucrose within the brain stem (Tokita &
Boughter Jr., 2016; Tokita et al., 2012). These investigators, however, did not screen
for glutamate-best neurons to determine if a similar convergence of sucrose signal-
ing on umami-best cells also occurs.
Currently, our understanding of neural processing of umami taste stimuli in the
NST and PBN in rodents is limited. For example, besides taste perceptual functions,
the NST and PBN are involved in post-ingestive effects of umami capable of direct-
ing behavior. However, little is known about how these structures contribute to post-
ingestive effects or if their perceptual and nonperceptual functions overlap. This
analysis may require experimentally distinguishing neural responses to umami,
sucrose, and NaCl to determine the presence or absence of glutamate-IMP syner-
gism. In addition, more precise analysis of the specific characteristics of taste-
evoked responses in the NST and PBN of awake and behaving animals, such as
those described by Roussin, D’Agostino et al. (Roussin et al., 2012), may be needed
to better understand how umami taste is distinguished from other taste stimuli in the
brain stem.
3.9 Thalamus
A dissociation between signaling of NaCl and MSG was reported for neuronal
responses recorded from the ventroposterior medial parvicellular (VPMpc) nucleus
of the thalamus when studied using amiloride to reduce the contribution from Na+
taste (Tokita & Boughter Jr., 2012). The addition of amiloride reduces the similarity
in response profiles of these neurons to NaCl and MSG but has no effect on the rela-
tively weak correlations between MSG and other basic tastes such as sweet
(Verhagen et al., 2005). Thus, the greater impact of amiloride on NaCl responses than
on MSG responses suggests that glutamate signaling may follow a channel separate
from that for sodium. Neural fMRI (functional magnetic resonance imaging) data
also suggest that umami and salty taste sensations are processed somewhat differ-
ently in the thalamus of humans (Iannilli et al., 2012; Han et al., 2018). Whether
such differences between sweet and glutamate signaling also exist has not yet been
adequately tested.
3.10 Forebrain
Responses of the FOI, the primary gustatory cortex (see Fig. 3.3), to umami stimuli
are of particular interest, because generally this is considered where quality-intensity
discriminations are made, at least in monkeys and humans. Much of the earlier work
on cortical responses to umami was in nonhuman primates, primarily macaque
monkeys (Scott et al., 1986; Scott & Plata-Salaman, 1999; Scott et al., 2001). Baylis
and Rolls (1991) reported finding neurons in the macaque primary taste cortex and
caudolateral OFC (a secondary taste cortex) that responded best to glutamate. These
glutamate-best neurons were of approximately the same number and exhibited simi-
lar responsiveness to glutamate as neurons tuned to respond to glucose or any of the
other basic tastes. Moreover, responses in these glutamate-best cells did not corre-
late well with responses to NaCl or sucrose. In the macaque caudolateral OFC,
cortical cells exhibited response profiles for MSG independent of NaCl or any of the
other basic tastes (Baylis & Rolls, 1991; Rolls & Baylis, 1994). Moreover, evalua-
tion of the reward value and pleasantness of umami stimuli appears to occur in
the OFC.
In rats and mice, recent studies of gustatory cortex have capitalized on innovative
methods to relate neural responding with behavior. A two-photon imaging study
detected discrete areas within insula layers 2 and 3 that responded to discrete stim-
uli, including umami (Chen et al., 2011). Stapleton et al. (2002), using temporal
assays of cortical responses to taste stimuli with multielectrode arrays of gustatory
cortex while a rat performed a simple taste discrimination, found that individual
cortical neurons responded to MSG stimulation with action potential patterns dis-
cernable from responses to sucrose, NaCl, or other stimuli. Moreover, in some
cases, the responses to these stimuli were in the opposite direction. For example,
even though a cortical cell increased its firing rate to increasing concentrations of
MSG, the same cell could show a decrease in response to increasing concentrations
of sucrose. Similar temporal analyses of gustatory nerves and brain stem structures
may reveal further differences between umami taste signaling and other basic tastes
in the rodent.
In humans, fMRI has also revealed that umami stimuli can activate unique areas
of the human FOI, as well as areas shared with other basic tastes. In studies compar-
ing MSG with NaCl and other taste stimuli, significantly different activation pat-
terns in the FOI were evoked by umami, NaCl, and sucrose stimuli (Han et al., 2018;
Singh et al., 2011; Prinster et al., 2017). De Araujo et al. (2003) found activation of
the rostral FOI, the caudolateral OFC, and the rostral anterior CG by taste stimula-
tion with 1 M glucose, 0.05 M MSG, 0.005 M IMP, or the combination of MSG and
IMP. Careful analysis of a 30-voxel area of the left OFC showed evidence of activa-
tion by the MSG-IMP mixture consistent with synergy between the two umami
substances (de Araujo et al., 2003). In a follow-up study, McCabe and Rolls (2007)
examined fMRI activation with 0.1 M MSG and a savory vegetable odor presented
individually or as a mixture. Subjects subjectively rated the pleasantness of the
MSG-odor combination as greater than MSG alone. Cortical activation by the
combination was significantly greater in the medial OFC and the pregenual CG than
expected by the summed activation of the individual stimuli and correlated with
pleasantness ratings by individual subjects. Importantly, these data illustrate how
glutamate can increase the palatability of a food when combined with a consonant,
savory odor (Rolls, 2009).
Neuroimaging studies have also given us insights into cortical control over higher-
order or “top-down” cognitive functions on the perception of umami. Secondary
taste cortices such as areas of the prefrontal cortex and the CG are the main regions
involved in these functions, especially as they affect the pleasantness of umami stim-
uli. As predicted from monkey electrophysiological research described above, fMRI
studies have shown that the response of the human OFC to umami stimuli decreases
with satiation, an effect not seen in the FOI (Luo et al., 2013). In addition, an area of
the OFC exhibits synergistic activation to the combination of MSG and IMP (de
Araujo et al., 2003) and receives input from the olfactory system (McCabe & Rolls,
2007). Collectively, these findings indicate the OFC is strongly involved in determin-
ing the perceived pleasantness and flavor of taste stimuli.
Cognitive modulation of pleasantness is mediated by other areas of the brain as
well. For example, the affective dimension of the pleasantness of umami appears to
activate areas of the pregenual CG and the ventral striatum, areas that receive input
from the OFC (Grabenhorst et al., 2008). Moreover, the degree of activation of these
areas and the behavioral responses associated with the affective property of umami
can be modulated by word labels. Depending on the nature of the task, attentional
processes can selectively enhance activation of these areas (Grabenhorst et al.,
2008). For example, activation of OFC, but not the FOI, is increased when the task
focuses on the pleasantness of umami. However, if the task focuses on evaluating
the intensity of umami stimuli, activation of the FOI, but not of the OFC, is increased
(Grabenhorst et al., 2008). Understanding how umami affects cognitive processes
may have important clinical implications (Magerowski et al., 2018). When umami
is added to food items, subjects increase their preference for and intake of these
foods (Bellisle, 1998, 1999). This information could help patients with dietary chal-
lenges, such as the elderly, those affected by cardiovascular disease, or those with
taste deficits from chemotherapy or toxic agents.
3.11 Umami Signaling in the Gut: Gastrointestinal System
In one sense, the gastrointestinal (GI) tract can be viewed as a long, convoluted
tubular chemosensing structure with different chambers specialized for digestion
and absorption. Glutamate sensing in the oral cavity activates the cephalic phase of
digestion, but glutamate is sensed again in the gut, which enhances digestive pro-
cesses (vago-vagal reflex) and influences cognitive processes related to umami per-
ception via the gut-brain axis. Throughout the GI tract, enterochromaffin sensory
cells detect the chemical composition of ingested food and chyme. These enteric
endocrine cells secrete serotonin and certain gut hormones, including
cholecystokinin, gastric inhibitory peptide, glucagon-like peptides (GLP-1, GLP-2),

peptide YY, and others. The enteroendocrine cells have different names (e.g., endo-
crine I cells and endocrine L cells) depending on the peptide they secrete.
Vagal afferents do not directly innervate gut sensory cells but, rather, are acti-
vated by paracrine hormonal signals released by enteroendocrine sensory cells typi-
cally expressing a receptor also found in the oral cavity (Akiba & Kaunitz, 2011;
Raka et al., 2019). For example, the metabotropic glutamate receptor mGluR1, ini-
tially found in the oral cavity, is also expressed in certain gut neuroendocrine cells
(San Gabriel et al., 2005, 2007; Nakamura et al., 2010; San Gabriel & Uneyama,
2013). L cells also express receptors found in the oral cavity, such as T1R, T2R, and
calcium-sensing receptor (CaSR) families capable of detecting sweet, umami (and
other amino acids), and bitter compounds (Uematsu et al., 2011; Raka et al., 2019).
Similar to taste cells in taste buds, these cells also have GPCR proteins and signal-
ing pathways. When activated, L cells release GLP-1 and GLP-2. I cells express
T1R1/T1R3 and CaSR receptors, which when activated release cholecystokinin.
These peptides activate other enteroendocrine cells and vagal afferents. Abdominal
vagal innervation extends from the esophagus to the upper GI tract and serves as the
primary neuroanatomical component of the gut-brain axis. It relays information
about gut content to the brain, which can modulate GI functions (e.g., digestion,
absorption, emptying) and conscious sensations (e.g., satiety, taste perceptions)
(Tome, 2018). Intragastric loading studies typically show that gut sensing of
ingested substances either adds to or subtracts from signaling of the oral pathways.
Postingestive effects of umami stimuli on taste perception appear to be quite
potent and more wide-spread than previously thought. Intragastric infusion with
MSG in mice and rats, when paired with an aversive agent, can lead to learned
avoidance of glutamate or, if paired with a flavor, can enhance flavor preferences
(Ackroff & Sclafani, 2016). Although the associative processes underlying these
effects are not known, fMRI studies in mice detected neural activation in the dorsal
vagal nucleus, the NST, and the insular cortex following GI infusion of glutamate.
GI activation of these areas can be combined with activity induced by oral sensa-
tions and the lateral hypothalamus and thereby influence cortical regulation of eat-
ing behaviors. This activation is reduced by vagal nerve cut and is abolished by a
variety of serotonin inhibitors and by a nitric oxidase inhibitor, suggesting this sig-
nal is mediated by serotonin and nitrous oxide (Tsurugizawa et al., 2009, 2010;
Uematsu et al., 2010, 2011; Torii et al., 2013). In humans, the postoral ingestive
effects of MSG and other taste compounds were examined using a naso-oral tube to
bypass the oral cavity during a memory task (Meyer-Gerspach et al., 2016). fMRI
revealed that in the sessions in which MSG was administered, more activation was
observed in FOI areas (primary taste cortex), the CG, Brodmann’s area 7, and pre-
cuneus cortical areas (associated with emotional, mnemonic, and conscious infor-
mational processing of taste stimuli) than with sucrose or NaCl. These results
suggest that MSG may have stronger effects on areas involved in working memory
than seen with other taste compounds. It is unclear if these effects are comparable
to those found in rodents, but they suggest that glutamate and the gut-brain axis may
play a larger role in cognitive processing than previously suspected.
3.12 Summary and Conclusions
The savory taste and mouthfeel of umami compounds, notably MSG, are generated
by receptor cells and neurons of the gustatory sensory system, complemented by
inputs from cells lining the GI tract. The existence of specialized GPCRs unique for
umami compounds (T1R1 + T1R3, taste-mGluR1, taste-mGluR4) in taste buds and
GI tract cells reinforces the notion that umami is indeed a basic taste alongside
sweet, sour, salty, and bitter. Neuronal responses to umami compounds at all levels
of the gustatory system in the CNS and peripheral nervous system often overlap
somewhat with responses to NaCl (salty) and sucrose (sweet). This suggests that the
neural circuitry for umami, sweet, and salty taste may partially overlap. Nonetheless,
there is substantial evidence that substances that elicit an umami taste generate
afferent signals that are both complex and unique and that these signals are the basis
for differential processing of umami taste in rats, mice, nonhuman primates,
and humans.
To date, the focus of much of umami research has been to determine if umami
taste is worthy of the status of a basic taste. However, this may well have caused
researchers to ignore a more complex and quite possibly much more significant
question: how do glutamate, IMP, and other umami stimuli affect the taste of other
substances? The interactive nature of the community of cells within a taste bud is
just now becoming apparent (Roper & Chaudhari, 2017; Rodriguez et al., 2021) and
may play an important role in umami-related enhancement of taste signaling within
the oral cavity. However, the overlap of umami, salt, and sweet neural pathways, a
feature of the CNS taste system that has made it so difficult to find umami-best
neurons, may be key to umami’s ability to interact with other tastes. A reasonable
and testable hypothesis is that umami signaling can modify neural signals generated
by complex taste mixtures and natural stimuli at one or more levels of the CNS. If
so, then the challenge is to determine how the umami signal interacts with other
taste signals within these CNS structures to modify taste perception.
At least two directions suggest themselves as fruitful starting points to explore
umami taste processing in the brain. One approach would be to investigate the tem-
poral pattern of taste-evoked neural responses (“taste code”) elicited by the interac-
tion of MSG/IMP and other taste stimuli at the several levels of gustatory signal
processing in the brain, perhaps through ensembles of neurons in these overlapping
pathways (e.g., Stapleton et al., 2006; Katz et al., 2002; Di Lorenzo & Victor, 2003;
Di Lorenzo et al., 2009; Roussin et al., 2012; Sammons et al., 2016). A second
approach would be to use natural foods rich in umami as gustatory stimuli and
investigate how signals generated by these stimuli are processed at all levels in the
gustatory nervous system, from taste buds to the cortex (e.g., Delay & Kondoh,
2015; Sammons et al., 2016; Pilato & Di Lorenzo, 2018). Studies such as the above
not only would reveal important information about the basic physiology of umami
taste but also would increase our understanding of how umami might be utilized
with human populations—such as the elderly or patients with dietary issues—to
improve nutritional intake.
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Dr. Eugene R. Delay earned a BS Degree in Psychology at the University of Idaho in 1972.
While at the University of Georgia, he earned an MS Degree in 1977 and a PhD in 1979 in
Biological Psychology. He held faculty positions at Regis University (Denver, CO) and the
University of Vermont (Burlington, VT). His research career began in psychopharmacology before
shifting focus to recovery of function after brain injury. In 1991, he began a long-standing collabo-
ration with Dr. Roper examining receptor and transduction mechanisms underlying umami taste.
Recently, he studied the effects of chemotherapy drugs on the taste system, including umami.
Stephen D. Roper received a Bachelor of Science Degree at Harvard College in 1967, his PhD
from University College London in 1970, and a postdoctoral training at Harvard Medical School
until 1973. Dr. Roper then held faculty positions at Harvard Medical School, University of
Colorado Medical School, Colorado State University (where he was Chairman of Anatomy and
Neurobiology), and presently Miller School of Medicine, University of Miami. In 1996, Drs.
Roper, Delay, and N. Chaudhari identified the first taste receptor, the umami receptor, and subse-
quently have investigated the molecular, cellular, and behavioral aspects of umami taste.
Chapter 4
Umami and Salty: A Cooperative Pair
Aubrey Dunteman and Soo-Yeun Lee
4.1 Introduction
A specific preference for substances exhibiting a salty taste, typically originating

from foods containing salt (NaCl), has been identified in humans. Salt is used in
food for a variety of purposes, largely falling into the categories of preservation,
flavor, and processing function. While salt is useful in food products, when con-
sumed in excess, it may increase the risk of many chronic diseases, such as cardio-
vascular disease and hypertension. In attempts to reduce the high-sodium content of
many processed and prepackaged foods, the addition of umami flavor has been
increasingly investigated due to its potential to enhance saltiness, as well as other
favorable flavor attributes. Much investigation has focused on glutamates, primarily
monosodium glutamate (MSG), although other frequently studied umami sub-
stances include soy and yeast derivatives. The many options available to impart
umami taste have varying results, generally positive overall. Despite the variety of
investigations into using umami to reduce sodium in food, there are several gaps in
the research that would benefit from future study. Examples include how to enhance
umami in solid food matrices aligned with sodium reduction and how greater reduc-
tions in sodium content can be achieved by combining umami enhancement and
physical modification methods.
A. Dunteman · S.-Y. Lee (*)

University of Illinois, Urbana, IL, USA
e-mail: [email protected]; [email protected]

https://doi.org/10.1007/978-3-031-32692-9_4
74 A. Dunteman and S.-Y. Lee
4.2 Role of Salt in Food
4.2.1 Relevance of Salt
It has become commonplace that, when food is bland or unappetizing, salt is added
in an attempt to improve flavor. Given the ubiquitous presence of salt and its exten-
sive use in various cuisines, it comes as no surprise that humans have a specific
preference for substances with a salty taste. Whether this preference originates from
an innate appetite for salt or is largely a learned behavior remains unclear (Círillo
et al., 1994; Mennella, 2014). The affinity for salty foods may also be partially
explained by the biological necessity of sodium in the diet for survival, which is
most easily accessible through sodium chloride, popularly known as salt (Beauchamp
& Engelman, 1991; Liem, 2017). Regardless, the inclination for humans to con-
sume salty foods is apparent, as demonstrated by the preference for salted over
unsalted foods and the prevalence of sodium overconsumption (Beauchamp &
Engelman, 1991; Verma etal., 2007).
4.2.2 Major Roles of Salt
Extending past salt appetite, dietary sodium intake is also driven by functional roles
of salt (hereafter referred to as NaCl) in food. By grouping certain roles together,
three main uses of NaCl can be observed: preservation, processability, and flavor
(Kilcast & den Ridder, 2007) (Fig. 4.1).
4.2.2.1 Preservation
NaCl has been used throughout history as a preservative, with many traditional fer-
mented, brined, and pickled products relying on its ability to preserve foods (Everis
& Betts, 2019; Kim et al., 2017). While NaCl itself does not confer significant
Fig. 4.1 Main roles of NaCl used in food products. (Created with BioRender.com)
4 Umami and Salty: A Cooperative Pair 75
antimicrobial activity, its ability to create an inhospitable environment for microor-

ganisms and pathogens effectively extends a product’s shelf life by preventing rapid
spoilage. NaCl confers stress on microbes in part by reducing water activity, which
limits unbound water available for microbial growth, and by increasing osmotic
pressure, which can inhibit microbial growth and survival by inducing osmosis
(Albarracín et al., 2011; Henney et al., 2010; Kim et al., 2017). These mechanisms
are pivotal to interrupting harmful microbial processes; hence, NaCl is an easily
accessible and inexpensive method of preservation.
4.2.2.2 Processability
Another major role of NaCl in foods relates to its functionality during the process-
ing of certain food products. For example, in the manufacturing of bread, NaCl is
necessary for adequate gluten development in dough, due to its ability to strengthen
gluten, increase dough elasticity, and decrease dough extensibility (Noort et al.,
2012; Simsek & Martinez, 2016). The presence of NaCl also impacts the fermenta-
tion process: by increasing NaCl levels in dough, yeast activity can be reduced, and
as a result, the rate of fermentation decreases (Cauvain, 1998). NaCl is also impor-
tant for the production of certain meat products, influencing water-holding capacity,
activating enzymatic activity, and potentially generating ripening pigments
(Albarracín et al., 2011). Similarly, NaCl is added to cheese to alter the water-
binding activity of casein, to modify protein conformation by controlling certain
enzymatic activity, and to regulate the growth of the lactic acid bacteria used in
production (Albarracín et al., 2011; Floury et al., 2009). The examples listed above
may also be relevant to the production of other products as well, and the functional
role that NaCl plays in the processing of food is not limited to these examples.
4.2.2.3 Salty Taste
The effect that NaCl has on flavor is likely its most recognizable role in food.
Evident from the name of the taste, salt confers a salty taste, one of the five identi-
fied basic tastes alongside sweet, sour, bitter, and umami, and has been found to
have flavor-enhancing qualities. When NaCl is omitted from certain foods, the fla-
vor quality is often found to suffer. Resulting food products may be described as
tasteless, bland, or insipid (Cauvain, 1998; Kilcast & den Ridder, 2007). To reduce
sodium levels in food, other chloride salts have been suggested in anticipation that
they may also impart the easily identifiable salty taste. While many chloride salts
exist and have been used in food manufacturing, only NaCl and LiCl have been
found to impart a salty taste (Van Der Klaauw & Smith, 1995). With a goal of reduc-
ing sodium, LiCl appears to be an appropriate replacement for NaCl as it imparts
saltiness in a manner similar to sodium’s ion channel utilized with NaCl. However,
due to the toxicity of lithium ions, LiCl has been eliminated as a possible replace-
ment (McCaughey, 2019).
Although NaCl’s use to impart a salty taste has not been successfully replicated,
other compounds yet to be investigated likely have the potential to add saltiness to
food. Looking ahead, it is sensible to consider that the activation of the sodium
receptor mechanism may not be the sole determinant of salty taste perception—we
should not assume there is only a single receptor, with only one transduction cas-
cade, for such a basic taste (McCaughey, 2019). Considering how taste cells also
contain separate sweet- and umami-responsive receptors or pathways that are inde-
pendent of the canonical T1R3 receptor (Damak et al., 2003), the salty taste mecha-
nism may also involve multiple receptors and has yet to be completely
characterized.
4.2.2.4 Flavor Modification
Although NaCl is best known for conferring a salty taste, its influence on flavor
extends past that singular taste modality. The sodium in NaCl can act as a bitter
blocker and can enhance sweet tastes. The ability for sodium and NaCl to suppress
bitter taste has been demonstrated alongside such bitter compounds as quinine
hydrochloride (Schifferstein & Frijters, 1992) and caffeine, magnesium sulfate,
amiloride, potassium chloride, and urea (Beauchamp et al., 2001; Breslin &
Beauchamp, 1995), to various percentages of maximum bitterness sensation. This
suppression of bitterness is likely a result of suppression at the periphery, in the oral
cavity, where tastes interact with receptor cells and where, consequently, sodium
and NaCl can inhibit taste receptor function (Wilkie et al., 2014). This is supported
by the ability of sodium and NaCl to suppress bitterness when there is minimal salty
taste perception (Keast & Breslin, 2002), but not when bitter and salty stimuli are
applied to opposite sides of the tongue simultaneously (Kroeze & Bartoshuk, 1985).
In contrast to evidence of NaCl acting as an effective bitter blocker, some research
has reported sodium or NaCl to be ineffective at reducing caffeine bitterness percep-
tion (Kamen et al., 1961), possibly suggesting that the suppression effect depends
on such factors as the specific sodium compound and its concentration and the bitter
compound and its concentration. Similarly, sodium’s or NaCl’s effectiveness may
be affected by the consumer’s taste phenotype: those who perceive 6-n-
propylthiouracil (PROP) as more bitter experienced bitter blocking from sodium
compounds, when tested with Brassicaceae vegetable, to a greater extent than those
unable to taste PROP at all (Sharafi et al., 2013).
The ability for sodium or NaCl to suppress bitterness may also enhance favorable
flavors by releasing tastes from the mixture suppression effect: the suppression of
bitter tastes by sodium salts releases sweet tastes from being suppressed by bitter-
ness (Beauchamp et al., 2001; Breslin & Beauchamp, 1997; Kemp & Beauchamp,
1994). The phenomenon can be particularly useful for increasing vegetable intake,
as the increased sweetness and decreased bitterness may increase consumer liking
(Sharafi et al., 2013; Wilkie et al., 2014).
4.3 Sodium Intake and Health
4.3.1 State of Sodium Consumption
The overconsumption of sodium, well above recommended intake limits, has

become commonplace worldwide (Brown et al., 2009). Despite the recommenda-
tion by the Dietary Guidelines for Americans for sodium intake to be limited to less
than 2300 mg per day (U.S. Department of Agriculture and U.S. Department of
Health and Human Services, 2020), and the World Health Organization’s recom-
mendation of less than 2000 mg/day (Brown et al., 2009), the average sodium intake
for American adults is roughly 3400 mg/day; thus, sodium has been designated as a
nutrient of health concern for overconsumption (Dietary Guidelines Advisory
Committee, 2020).
4.3.2 Sodium Sources
An estimated 75% of sodium intake in North American and European diets has been
found to originate from processed or restaurant foods; roughly 10–12% can be
traced each to sodium occurring naturally in foods or from discretionary use at the
table (James et al., 1987; Mattes & Donnelly, 1991). Most sodium intake in devel-
oped countries results from consumption of “hidden” sodium in processed foods.
However, some countries also struggle with high-sodium intake from different ori-
gins. For example, in the People’s Republic of China, over 75% of dietary sodium
has been found to come from salt added in home cooking (Anderson et al., 2010;
Zhai, 2006).
4.3.3 Sodium’s Effects on Health
Evidence of associations between excess sodium consumption and a multitude of

adverse health effects has been demonstrated by meta-analyses and systematic
reviews (Institute of Medicine, 2005; Malta et al., 2018). Figure 4.2 displays a
selection of these health effects. A growing body of literature on dietary sodium
intake and health has found a linear relationship between sodium intake and risk of
developing hypertension (Bock & Cottier, 1961; Hermansen, 2000; Kim & Andrade,
2016), which further increases the risk of developing such conditions as coronary
heart disease, congestive heart failure (Butler et al., 2015), stroke (Strazzullo et al.,
2009), and renal disease (Keiko, 2017). Research also suggests that excessive salt
intake leads to increased risk for certain cancers, including gastric cancer (Sheng
et al., 2012) and liver cancer (Sun et al., 2020), further demonstrating the need to
reduce the general population’s sodium intake.
Fig. 4.2 Some increased health risks from overconsumption of sodium. (Created with
BioRender.com)
4.4 Strategies to Reduce Sodium Intake
4.4.1 Stealth
Many methods have been investigated to reduce sodium overconsumption, such as

leading consumers to healthier choices and manipulating high-sodium foods to con-
tain reduced-sodium levels. While a sudden reduction in the amount of added salt in
a recipe can lead to lower acceptance by consumers, a stepwise reduction in
sodium—the stealth method—may alleviate some of the effects on consumer per-
ception without requiring additional ingredients. The stealth method can be effec-
tive in reducing sodium overconsumption by developing foods that remain liked by
consumers through progressively reducing the sodium content of the food or of a
consumer’s diet over time and, consequently, reducing preference for higher levels
of salt intensity. This shift in preference can become evident anywhere from a few
days to a few months after the shift in sodium intake (Bertino et al., 1982; Dötsch
et al., 2009; Girgis et al., 2003; Quilez & Salas-Salvado, 2012).
4.4.2 Physical Modification
Another method of reducing sodium in foods that can be used without including
additional ingredients is physical modification. Methods for physical modification
of foods for sodium reduction are diverse (Fig. 4.3) and include using different salt
crystal morphologies (Rama et al., 2013; Rodrigues et al., 2016), altering the texture
of a food (Pflaum et al., 2013), and inhomogeneous distribution of salt in a food
Fig. 4.3 Physical modification methods used to reduce sodium in food products. (Created with
BioRender.com)
product to produce taste contrast (Noort et al., 2010). While physical modification
methods have generally had promising results, costs associated with new or
upgraded machinery to produce those methods may be an obstacle for many manu-
facturing companies.
4.4.3 Flavor Modification
In flavor modification, the flavor of a food is modified by adding another compound

or food ingredient. Described below are only a subset of the wide range of flavor
modification strategies available, primary detailing salt replacement and umami-
based strategies.
4.4.3.1 Salt Replacement
The most widely investigated flavor modification method involves salt replacers.
Although many materials have been investigated for their potential as a salt replacer,
such as phosphates (Pandya et al., 2020; Seman et al., 1980), sulfates (Davaatseren
et al., 2014), and citrates (Braschi et al., 2009), the most promising are mineral salts,
in particular, potassium chloride (KCl) (Grummer et al., 2012). Although KCl can
contribute only slightly less saltiness than sodium chloride, it has also been com-
monly described to have a bitter taste, which further exemplifies the need for
improved methods for salt reduction (Rogério Tavares Filho et al., 2020). As noted
above, LiCl has been found to confer a salty taste similar to sodium chloride, but it
has been deemed unsafe to consume and thus is not a viable salt replacement option
(Hand et al., 1982).
4.4.3.2 Umami-Based Strategies
Fundamentals of umami Strategies
In 1985, a groundbreaking meeting to discuss umami was held. At this meeting,

umami was selected as the scientific term for the specific taste attributes generated
by glutamates, particularly monosodium glutamate (MSG), as well as the 5′-ribo-
nucleotides guanosine monophosphate (GMP) and inosine monophosphate (IMP)
(Tepper & Yeomans, 2017). Research has shown that significant differences exist
regarding individuals’ ability to detect MSG and that some may lack the ability to
detect MSG relative to NaCl (Lugaz et al., 2002; Raliou et al., 2009).
MSG is likely the most investigated umami-imparting substance for use in
sodium reduction. Despite the evidence of its safety (Fernstrom, 2007), stigma still
surrounds it from the “Chinese restaurant syndrome” it has been erroneously
accused of causing (Husarova & Ostatníková, 2013). While other glutamates similar
in structure to MSG, such as monoammonium glutamate (MAG), monomagnesium
di-L-glutamate (MDG), and calcium diglutamate (CDG) (see Fig. 4.4), have been
studied for their potential in maintaining acceptance or saltiness of reduced-sodium
foods, with some level of success, publications are limited (Ball et al., 2002; Carter
et al., 2011; Daget & Guion, 1989). IMP and GMP, as well as their respective
derived acid salts, disodium inosinate and disodium guanylate (Fig. 4.5), have also
been investigated for sodium reduction. Research has shown that they can be used
as an alternative to MSG, thus avoiding the stigma surrounding MSG. Disodium
inosinate and disodium guanylate are also used in a 50:50 mixture, referred to as I
+ G, which then may be used in conjunction with glutamates to impart a synergistic
effect and yield a highly savory taste with minimal concentrations. These nucleotide
compounds are produced by microbial fermentation, derived by animal origin or,
more often, all-vegetable tapioca starch, so their use may be more accepted by the
public than the well-known MSG.
Detection Thresholds of Umami Substances and Sodium Chloride
The detection thresholds of umami substances differ from that of sodium chloride
both in concentration and in threshold variability. The concentration required to
detect NaCl in water is lower than the concentration for MSG, first reported to be
1/400 and 1/3000, respectively (Ikeda, 2002). Studies on the interaction between
umami and the basic tastes at threshold have yielded conflicting results, although it
is more often reported that MSG does not change threshold values for sucrose or
sodium chloride (Lockhart & Gainer, 1950; Yamaguchi, 1998) but reduces thresh-
olds for sour and bitter tastes (Mosel & Kantrowitz, 1952). The impact of different
umami substances on detection thresholds of other compounds that elicit primarily
umami tastes, such as MSG or the 5′-ribonucleotides, is notable: both in clear soup
Fig. 4.4 Compound structures of glutamates: monosodium glutamate (a), calcium diglutamate
(b), monomagnesium di-L-glutamate (c), and monoammonium glutamate (d). (National Center
for Biotechnology Information, 2021a, b, c, d, e, f, g, h)
(Luscombe-Marsh et al., 2008) and in solution (Yamaguchi, 1998), when the sol-
vent contains IMP, the detection threshold for MSG is lowered significantly. This
also occurs with the IMP detection threshold when evaluated in an MSG-
supplemented solution, as MSG and IMP are known to have a strong synergistic
effect (Yamaguchi, 1998). When evaluated in solutions containing other basic taste
substances, the detection threshold of MSG did not increase except when sucrose
concentration was high. Because of the synergistic ability for the MSG detection
threshold to be reduced and the relative ease of distinguishing umami in the pres-
ence of other tastes, it has been suggested that, among the five basic tastes, MSG is
the most sensorily perceived substance (Yamaguchi, 1998). This is a point of con-
tention, though, as the intensity of umami sensation is considered inferior compared
to substances characterized by extremely strong tastes, including alkaloids or sac-
charin (Ikeda, 2002).
Fig. 4.5 Compound structures of ribonucleotides: disodium inosinate (a), inosine monophosphate
(b), disodium guanylate (c), and guanosine monophosphate (d). (National Center for Biotechnology
Information, 2021a, b, c, d, e, f, g, h)
Taste Profile of Umami Substances
Statements used to describe the taste and flavor of MSG vary greatly. From the first
symposium on MSG in 1948, many descriptions of pure MSG were reported,
including “a desirable meat-like taste,” “all four components: sweetness, sourness,
saltiness, and bitterness,” and “a persistent lingering flavor reaction,” to name a few
(Beauchamp, 2009). MSG has been described as having a slightly sweet and salty
taste (Lockhart & Gainer, 1950), but it has also been described as eliciting sweet,
sour, salty, and bitter tastes (Mosel & Kantrowitz, 1952). Despite numerous studies
investigating MSG taste, a clear consensus on its sensory properties is still lacking
(Kemp & Beauchamp, 1994; Beauchamp, 2009).
The temporal profile of tastes in solution varies depending on the substance and
whether it is alone or combined with other tastes. When evaluating a single primary
taste solution at a time, salty and umami tastes show similar temporal time-intensity
profiles and temporal dominance of sensations (Rocha et al., 2020). These two
tastes were also similar in that the duration of the taste solutions, MSG for umami
and NaCl for salty, increased in a concentration-dependent manner.
Saltiness of Umami Compounds
The saltiness of MSG is approximately 30% that of NaCl by molar sodium concen-
tration and 10% of NaCl by weight (De Souza et al., 2013; Yamaguchi, 1998). IMP
has a saltiness equivalence of approximately 50% molar sodium concentration of
NaCl and 7% by weight. Combining both MSG and IMP results in a taste that is
largely umami due to their synergistic actions and thus is negligible in saltiness
(Yamaguchi, 1998).
Interactions Between Salty and Umami Tastes
The effect of combining MSG and NaCl on saltiness has been examined over the
years. A compensatory relationship between the two has been suggested, where
more MSG is necessary as NaCl is reduced and vice versa. In soups, the optimal
combination of NaCl and MSG has been reported as approximately 2:1 ratio of
NaCl to MSG (Chi & Chen, 1992; Jinap et al., 2016; Yamaguchi & Takahashi,
1984), although the dynamics of palatability and saltiness vary for differing matri-
ces (Baryłko-Pikielna & Kostyra, 2007; Kim et al., 2014). It has also been demon-
strated that enhanced palatability from MSG largely depends on the presence of
NaCl, as MSG alone reduced hedonic perception in rice, whereas MSG plus NaCl
mitigated palatability loss from MSG (Yamaguchi, 1987; Yamaguchi &
Takahashi, 1984).
In NaCl and MSG mixtures, a temporal sequence of basic taste perception was
evident: the dominant taste prior to swallowing was saltiness, whereas umami was
the dominant taste after swallowing (Kawasaki et al., 2016). In NaCl and MSG
mixtures, umami duration in MSG solutions decreased with additional NaCl (Lioe
et al., 2005), while saltiness duration in NaCl solutions increased with additional
MSG (Kemp & Beauchamp, 1994). Such findings demonstrate a mixture-duration
suppression effect for NaCl in NaCl+MSG mixtures and that saltiness of concen-
trated NaCl has a greater impact on umami taste duration than on enhanced umami
taste (Kawasaki et al., 2016).
The ability for umami compounds to enhance salty taste in solution has also been
evident using the glutamate MAG and the nucleotides IMP and GMP. At every
NaCl concentration, saltiness enhancement was greater using MSG and MAG than
using either IMP or GMP, although all umami flavor enhancers displayed similar
sensory profiles (Rocha et al., 2020). Saltiness enhancement was also more apparent
when NaCl concentration was reduced in smaller amounts, indicating that these
substances may be most appropriately suited for foods requiring less drastic reduc-
tions in sodium content (Rocha et al., 2020).
4.4.3.3 Umami Compounds as Flavor Enhancers

in Sodium-Reduced Products
Glutamates
Liquids
Glutamates have frequently been evaluated for their potential in sodium-reduced
liquid foods over the years. In pumpkin soup, CDG and MSG have been demon-
strated to maintain or increase ratings in liking, flavor intensity, familiarity, and
richness under reduced-sodium conditions (Ball et al., 2002). Similar results were
demonstrated using CDG in sodium-reduced chicken broth, where liking and pleas-
antness were maintained (Carter et al., 2011). Chicken broth containing a variety of
different glutamates, including MSG, CDG, MDG, potassium diglutamate, and
ammonium glutamate, were investigated previously with varying levels of success.
All except ammonium glutamate and pure glutamic acid were preferred, according
to preference ranking (Daget & Guion, 1989).
MSG has been the most researched glutamate for sodium reduction in liquid
foods. In spicy soup, an approximate 32% reduction in sodium was feasible with
MSG replacing a portion of the salt (Jinap et al., 2016). Promising results for MSG
in sodium-reduced foods were also evident for tomato sauce (Rogério Tavares Filho
et al., 2020), clear soup (Yamaguchi & Takahashi, 1984), chicken soup (Wang et al.,
2019b), and certain vegetable soups (Roininen et al., 1996).
Semisolids
The use of MSG and KCl in margarine allows an approximate 47% reduction in
sodium content while maintaining salty taste and overall impression (Gonçalves
et al., 2017). When used in combination with KCl and either I + G or amino acids,
MSG successfully maintained liking at 50% and 75% salt reductions (dos Santos
et al., 2014). In mozzarella cheese supplemented with MSG and KCl, a 54% reduc-
tion in sodium was feasible while maintaining sensory quality (Rodrigues et al.,
2014). While KCl has been demonstrated to have a closer saltiness equivalence to
NaCl than does MSG, MSG does not enhance undesirable tastes in butter, as had
been evident with KCl (De Souza et al., 2013).
Solids
MSG has been demonstrated to enhance liking in reduced-sodium potato chips and
puffed rice snacks compared to their full-salt counterparts and to either maintain or
increase liking when consumers were informed of the reduction in sodium (Buechler
& Lee, 2019, 2020). The effect of informed versus blind tasting is not consistent,
however, as contrasting results were found in potato chips. In another study, the full-
salt treatment was least liked compared to the MSG treatments under blind condi-
tions, yet once participants were informed, the MSG treatments became less liked
(Kongstad & Giacalone, 2020), suggesting consumers may still have a bias against
certain flavor-enhancing compounds such as MSG. The use of MSG in reduced
sodium white and multigrain bread was found to have similar acceptability and
sensory characteristics when compared to their full-salt counterparts, although in
contrast to aforementioned studies, information about sodium reductions and MSG
inclusion had not impacted consumer perceptions (Dunteman & Lee, 2023a, b)
indicating the possibility that consumers are placing less importance on clean label-
ing of their foods than before.
5′-Ribonucleotides
Liquids
Although MSG is the most investigated umami-imparting substance for reducing
sodium, the use of 5′-ribonucleotides IMP and GMP, as well as their respective
derived acid salts, disodium inosinate and disodium guanylate, has also been the
focus of investigation. As noted above, disodium inosinate and disodium guanylate
are also used in the 50:50 mixture I + G, which then may be used in conjunction
with glutamates to yield a synergistic effect and impart a highly savory taste with
minimal concentrations.
Research has shown conflicting feasibility for use of 5′-ribonucleotides in liquid
applications. In vegetable soup at both low- and high-salt content, the inclusion of
an umami mixture consisting of IMP, GMP, and MSG (Roininen et al., 1996) evoked
increases in pleasantness, taste intensity, and ideal saltiness. Further evidence for
using these nucleotides was identified with chicken noodle soup: inclusion of a
mixture of MSG, IMP, and GMP led to increased intensity for such attributes as
overall flavor, umami taste, and mouthfeel (Leong et al., 2016). In contrast, when an
umami mixture of IMP, GMP, and KCl was incorporated into tomato sauce, the
flavor acceptance was significantly reduced compared to the control sauce. On the
other hand, when only KCl and IMP were added to the tomato sauce, there was no
reduction in the acceptance of the evaluated attributes (Rogério Tavares Filho et al.,
2020). The contrasting effectiveness of these nucleotides appears partially influ-
enced by the food application: while I + G in mushroom, red beet, and asparagus
soups contributed considerably to palatability enhancement, inclusion of MSG and
I + G in green pea cream soup largely incurred a negative effect on palatability
(Baryłko-Pikielna & Kostyra, 2007). Finally, IMP plus MSG enhanced perception
of savory taste and flavor intensity and increase consumer acceptance in reduced-
sodium chicken noodle soup (Miyaki et al., 2016).
Semisolids
When incorporated into a sodium reduction strategy for semisolid food matrices,
nucleotides and/or their derivatives appear to mitigate quality loss resulting from
sodium reduction or undesirable attributes characteristic of KCl in salt replacement.
Addition of KCl with IMP and GMP to sausage patties allowed up to 75% NaCl
replacement before acceptability was reduced (Pasin et al., 1989). In sausages con-
taining KCl as salt replacement, the addition of IMP and GMP with certain amino
acids led to higher quality ratings than for sausages with KCl alone and resulted in
no differences compared to the full-salt control (Campagnol et al., 2011a, b, 2012).
The addition of MSG to sausages reformulated with IMP, GMP, and certain
amino acids allows for further reductions in sodium content, masking undesirable
sensory changes resulting from replacing up to 75% of NaCl in the samples with
KCl and allowing for greater than 65% reduction of sodium (dos Santos et al.,
2014). The use of I + G and IMP alone with KCl salt replacement in reduced-
sodium tomato sauce was also effective in masking metallic notes (Rogério Tavares
Filho et al., 2020). The promising quality of GMP for masking negative attributes
from KCl has also been reported in nonmeat products, such as reduced-sodium
cheddar cheese. While cheeses with KCl and IMP were less accepted across all
attributes evaluated, those with KCl and GMP either maintained or increased attri-
bute acceptance compared to the control cheese (Grummer et al., 2013).
Solids
Little research has been done on solid foods using nucleotides as a strategy for
sodium reduction. One study reported potato chips and puffed rice snacks seasoned
with IMP, GMP, and MSG to have higher ratings of liking compared to the full-salt
control samples (Buechler & Lee, 2019). When consumers were informed of the
sodium reduction, the puffed rice snacks containing IMP, GMP, and MSG were
rated as the most liked, while potato chips maintained their liking ratings. Descriptive
analysis of the chips containing IMP, GMP, and MSG revealed that ratings for
umami aftertaste and meaty aftertaste were higher and ratings for salty aftertaste
were lower compared to the full-salt control chips. External preference mapping
indicated that this treatment of potato chips was well liked. In rice puff snacks, the
majority of evaluated attributes significantly differed across treatments, and external
preference mapping indicated that those seasoned with IMP, GMP, and MSG were
most liked (Buechler & Lee, 2020).
Other Amino Acids
Liquids
While research has been limited thus far on amino acids in liquid foods, those that
have been have not had a positive impact on reduced-sodium foods. Soup with glu-
tamic acid was characterized by a low overall quality, an acid taste, and a reduced
chicken flavor (Daget & Guion, 1989). Similarly, lysine used as a bitter blocker in
conjunction with KCl in reduced-sodium tomato sauce was ineffective: the treat-
ment sample had increased bitterness and metallic taste compared to the full-salt
control (Rogério Tavares Filho et al., 2020).
Semisolids
Glycine, lysine, and taurine have been evaluated in conjunction with salt replace-
ment and MSG- and nucleotide-supplemented semisolid food matrixes, with differ-
ing results. Glycine has been used in reduced-sodium frankfurters with acceptable
consumer perception (Wilailux et al., 2020). Lysine in reduced-sodium recon-
structed ham successfully maintained overall acceptability, appearance, taste, and
mouthfeel liking compared to a full-salt control (Guo et al., 2020). Lysine and tau-
rine in reduced-sodium fermented sausage, with 50% NaCl replaced with KCl and
supplemented with MSG, IMP, and GMP, produced suitable sensory qualities (dos
Santos et al., 2014). Lysine and taurine were also reported to increase taste accept-
ability in fermented sausages in which KCl replaced a portion of the NaCl; taurine
in particular had promising results, as all attributes evaluated maintained accept-
ability compared to the full-salt control (Campagnol et al., 2011a, b). Results were
similar with lysine addition: consumer acceptance of color, taste, aroma, and texture
in reduced-sodium sausages was maintained, although once 50% of NaCl was
replaced with KCl, certain sensory defects could not be mitigated (Campagnol
et al., 2012).
In cheese, amino acids were used in conjunction with other sodium reduction
methods, such as added umami substances, physical modification, and salt replace-
ment. Arginine has opposing results. Studies have shown reduced-sodium cheese
with KCl and arginine to be less liked than full-salt cheese without KCl or arginine
(Silva et al., 2017) and to have decreased saltiness and cheese aroma and increased
sour and bitter tastes (Silva et al., 2018). In contrast, reduced-sodium cheese con-
taining KCl and arginine had acceptable cheese flavor and overall acceptance com-
pared to reduced-sodium cheeses with KCl alone (Felicio et al., 2016). Substitution
of 35% salt with KCl in combination with glycine in rice porridge was feasible:
ratings for overall liking, overall flavor, saltiness, and other attributes did not differ
significantly from the full-salt control (Sriwattana et al., 2016).
Solids
At the time of writing, no research has been conducted on the use of amino acids to
reduce sodium in solid foods. This is a research area that needs to be explored in
the future.
Other Umami Substances

A great variety of ingredients with naturally occurring umami tastes have been iden-
tified. Select ingredients used for their umami-conferring properties in sodium-
reduced foods are displayed in Fig. 4.6.
Fig. 4.6 Umami-containing ingredients investigated to reduce sodium in food products. (Created
with BioRender.com)
Liquids
The use of umami substances in liquid foods to aid in maintaining quality alongside
sodium reduction has been investigated in several studies. Partial replacement of
salt with fish sauce, a natural source of glutamate and 5′-ribonucleotides, was dem-
onstrated to successfully maintain overall taste intensity and deliciousness in
chicken broth, tomato sauce, and coconut curry, at rates of 25%, 16%, and 10%
NaCl reduction, respectively (Huynh et al., 2016). Soy sauce, despite contributing
some sodium, has been found to allow for salt reductions of up to 50% in salad
dressing and between 17% and 33% in tomato soup without significant reductions
in overall taste intensity or pleasantness (Goh et al., 2011; Kremer et al., 2009).
Other research has also demonstrated the potential for salt replacement by soy sauce
for sodium reduction in tomato soup, with results suggesting levels of 24–33% to be
feasible (Kremer et al., 2013b).
Other umami substances investigated in liquid foods include yeast extract, mush-
room concentrate, and tomato concentrate. While not incorporating sodium reduc-
tion into their treatments, researchers have found that overall liking and overall
flavor liking in chicken soup increased with the addition of yeast extract and were
maintained with mushroom concentrate and tomato concentrate, thus suggesting the
investigated umami ingredients may be used to produce reduced-salt soups with
attributes deemed acceptable by consumers (Wang et al., 2019a).
Semisolids
A variety of different umami substances have been used in semisolid foods to aid in
sodium reduction, including, but not limited to, yeast derivatives, soy derivatives,
and ingredients naturally high in glutamates like tomatoes, mushrooms, seaweed,
and cheese. Yeast derivatives are often used in combination with KCl for a variety
of purposes, such as to reduce bitterness perception, increase saltiness intensity, and
increase consumer acceptance. Yeast extract in bread with partial salt replacement
by KCl allowed for 67% sodium reduction without affecting bread consumption or
causing sodium intake compensation with sandwich fillings (Bolhuis et al., 2011).
Fermented sausages also benefited from yeast extract to mitigate quality defects
from KCl and reduce sodium by 50% (Campagnol et al., 2011a, b). Similarly,
Debaryomyces hansenii yeast inoculation improved taste quality of 17–20% salt-
reduced dry fermented sausages (Corral et al., 2014). Reduced-sodium prato cheese
with KCl salt replacement and supplementation with yeast extract and Lactobacillus
casei increased consumer liking and saltiness intensity and decreased bitterness
intensity and bitterness aftertaste compared to the same treatment without yeast
extract, to the same ratings as the control, allowing for an approximate 35% reduc-
tion in sodium content (Silva et al., 2017, 2018).
Soy derivatives have been used for sodium reduction in semisolid foods such as
bread and stir-fried pork. Over an exposure period of 15 days, liking of reduced-
sodium bread with soy sauce did not decrease, whereas liking of the full-sodium
control bread decreased; however, no differences were found in perceived saltiness
intensity (Kremer et al., 2013a, b). In stir-fried pork with soy sauce, a 29% reduc-
tion in sodium content was feasible without significant losses in ratings of product
pleasantness or overall taste intensity (Goh et al., 2011; Kremer et al., 2009).
Frankfurters have also benefited from soy sauce, allowing for a 20% reduction in
salt content without reduced ratings for quality or sensory characteristics; when
treatments included KCl in addition to the soy sauce, a 35% reduction was feasible
(McGough et al., 2012a, b). Additionally, no differences in overall liking or salti-
ness liking were observed for bacon, beef jerky, or ham with a 30% reduction of
sodium and incorporation of KCl and soy sauce (Shazer et al., 2015).
Mushrooms have been used in many studies of reduced-sodium semisolid foods,
although their purpose tends to focus on replacing other meats. In carne asada, sub-
stitution of a portion of beef with mushroom had no effect on flavor intensity, and in
beef taco blends, incorporating 50–80% ground mushroom increased overall flavor
intensity. Despite this, inclusion of mushroom did not fully mitigate saltiness reduc-
tion in the reduced-sodium blend (Myrdal Miller et al., 2014), but when 20% of beef
was replaced with mushroom, overall liking was maintained with a 25% reduction
in salt (Guinard et al., 2016). On a similar note, consumers preferred the reduced-
sodium filling with 45% mushroom over both the full-sodium control with all meat
and the full-sodium control with 45% mushroom (Wong et al., 2017). Mushrooms
were also successfully used in reduced-sodium beef patties at 20% replacement
levels: ratings for overall liking and saltiness liking were maintained with an
approximate 25% reduction in sodium (Wong et al., 2019). Mushroom extract has
also been investigated for sodium reduction in beef patties, where a 50% reduction
in salt in the presence of the 20% mushroom homogenate extract increased accep-
tance of a variety of sensory attributes and enhanced salt perception to levels similar
to that of the full-salt control and to levels greater compared to the 50% reduced-
sodium treatments with 5% or 12.5% mushroom homogenate extract and the 75%
reduced-sodium treatments (Mattar et al., 2018).
Other umami ingredients such as tomatoes, cheeses (Dos Santos et al., 2020;
Xiang et al., 2017), fish sauce (Huynh et al., 2016), hydrolyzed vegetable proteins
(Khetra et al., 2019), and seaweed derivatives (Barbieri et al., 2016; Vilar et al.,
2020) have also been investigated for use in reduced-sodium foods, with mixed
results, although results trended toward acceptable products by either maintaining
or increasing quality attributes.
Solids
At the time of writing, no research has been conducted on the use of umami sub-
stances to reduce sodium in solid foods. This is another research area in need of
exploration.
4.5 Conclusion
This chapter has discussed certain gaps in knowledge surrounding the use of the
umami taste in relation to salt and sodium reduction. While glutamates, particularly
MSG, are well studied, there is limited research into other approaches that would
align with consumer expectations. Soy and yeast derivatives, ingredients naturally
high in glutamates, and potentially certain amino acids may all be preferred by the
general public compared to glutamates or the 5′-ribonucleotides, given the familiar-
ity and perception consumers likely have surrounding these substances.
Research including both blind and informed conditions on umami-based sodium
reduction methods is largely absent from the literature yet would provide valuable
insights into the potential of each method once available on the market. There is also
a large gap in how these methods may fare in solid food products—studies currently
are limited to potato chips and puffed rice snacks. While this may reflect the fact
that most high-sodium foods are either liquids (e.g., soups) or semisolids (e.g.,
cured meats), further investigation into other product categories would be valuable,
as future food trends cannot be predicted. Research on umami in solid foods would
also help increase our knowledge of sodium reduction methods in solid foods.
Lastly, while much research has used umami-based methods in combination with
other umami substances or with the salt replacer KCl, other sodium reduction meth-
ods achieved by modifying the physical form or processing conditions in conjunc-
tion with umami taste have not been well studied; this approach may enable larger
reductions in sodium content than currently available.
Despite the diversity in sources of umami taste and sodium reduction presented
throughout this chapter, certain limitations have arisen during writing. Not every
piece of literature investigating the umami taste has been reviewed; gray literature
that is difficult to identify may be present yet inaccessible. Similarly, not every lit-
erature review on the topic has been included owing to limitations in space and in
the scope of this book. Although this chapter provides a detailed look into how
umami and salt are related, further connections may have not been identified as a
result of the tendency to present each taste independent of the others, without suf-
ficient literature review and discussion.
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Aubrey Dunteman, Ph.D. is a sensory scientist and recent graduate of the Department of Food
Science and Human Nutrition at the University of Illinois. Her graduate research focused on strate-
gies to maintain the palatability of reduced-sodium foods with a particular interest on the effect
that the umami taste can have on saltiness perception. She has published multiple review articles
on the outcomes of various sodium reduction strategies. Aubrey received her bachelor’s degree
from the University of Arkansas.
Soo-Yeun Lee, Ph.D. Dr. Lee is a Professor and Director of the School of Food Science at the
Washington State University. She is an accomplished scholar whose expertise is in sensory sci-
ence. Her research focuses on utilizing sensory evaluation to develop health-targeted new product
alternatives to promote healthful eating habits. She has over 100 publications and garnered about
$10 Million as PI and co-PI. Soo has also been recognized as an educator with many national
teaching awards, such as the IFT Cruess Award. Dr. Lee received her doctorate from UC Davis and
her bachelor’s degree from Yonsei University in Korea.
Chapter 5
Umami and Satiety
Martin R. Yeomans
Umami is today recognized as one of the five basic tastes, complementing the more
widely known salty, sweet, sour, and bitter tastes. In all five cases, the taste percept
(i.e., the conscious experience of the taste quality) arises from oral detection of
nonvolatile chemicals dissolved in saliva that bind to specific taste receptors on the
tongue. An important feature of all five basic tastes is that their qualities are unique:
that is, it is not possible to generate a different taste quality by mixing other tastes
(perceptual independence). In the case of umami, the primary receptor is known to
be a heterodimer of two proteins, TAS1R1 + TAS1R3, that has a unique role in taste
sensing (Damak et al., 2003; Li et al., 2002), and recent studies suggest strong posi-
tive selective pressure for the genes for both TAS1R1 and TAS1R3 in human evolu-
tion (Valente et al., 2018). These same receptors are found in other parts of the
digestive tract, but only those in the mouth give rise to the perceptual experience of
umami taste. A key question, then, is why we have evolved the ability to detect
umami? To understand this, we first need to consider the potential evolutionary
advantage of umami sensing.
Evolution shapes physiology to adapt to specific challenges for survival. For
omnivores, including humans (as cogently argued as the “omnivore’s paradox” in
(Rozin, 1976)), the key problem is how to identify which of the many potential
foods that occur in nature are safe and nutritious. On that basis, it has long been
argued that our ability to taste certain compounds in potential foods evolved to meet
that challenge, with the mouth the gatekeeper that directs decisions on whether a
potential food is ingested or rejected (Tanaka et al., 2007; Kim et al., 2004), inter-
preting taste as the “nutritional sense” (Bartoshuk, 2018). Accordingly, our ability
to sense bitterness ensures we are cautious when ingesting substances that could be
harmful (Shi et al., 2003; Breslin, 2013), and there is clear evidence of how diet
M. R. Yeomans (*)
University of Sussex, Brighton, UK

https://doi.org/10.1007/978-3-031-32692-9_5
102 M. R. Yeomans
shaped the evolution of bitter taste perception (Li and Zhang, 2014). Similarly,
sweetness may signal a safe source of energy (Beauchamp, 2016), and most recently
sourness has been proposed to help identify key vitamins in states of vitamin defi-
ciency (Teng et al., 2019). From this evolutionary perspective, what may be the
reason that humans can detect umami taste? The answer, based on the nature of the
foods that contain the compounds that generate umami taste, is that umami may
have evolved to help us sense sources of protein, an idea dating back to the original
discovery of umami taste (Ikeda, 1908) and then developed by other researchers
(e.g., Naim et al., 1991; Kurihara, 2009; Geraedts et al., 2011). But our ability to
detect umami compounds in the mouth and later in the gut could have a function
beyond the simple sensing of the presence of protein. Sensors in the mouth and gut
are also key components of appetite control, and recent evidence suggests that
umami taste not only allows us to sense the likely presence of protein but also helps
the body regulate protein intake. This chapter evaluates current state of play for
research that has tested the role of umami in satiety.
5.1 Why We Taste Umami: Lessons from the Diet
The original idea of umami relating to protein sensing was based on consideration
of where in the diet we find the chemicals that activate umami taste receptors.
Umami taste can be traced back to the initial isolation of glutamic acid from sea-
weed and the subsequent discovery in 1908 that it, and particularly its sodium salt,
monosodium glutamate (MSG), conveys a unique “flavor.” MSG is now known to
be the main molecule that stimulates human umami taste receptors. Subsequent
research found that the umami taste generated by MSG was greatly potentiated by
the presence of ribonucleotides, most specifically by disodium 5′-inosine mono-
phosphate (IMP) and disodium 5′-guanosine monophosphate (GMP) (Kuninaka,
1981; Fuke & Ueda, 1996). Notably, neither IMP nor GMP elicits a strong umami
taste alone, but both act synergistically with MSG to elicit a stronger umami taste
than predicted by the sum of the individual components (Yamaguchi & Ninomiya,
2000; Fuke & Ueda, 1996). To understand why we have evolved the ability to taste
umami, we need to consider where in our food supply we find MSG, IMP, and
GMP: in a wide range of foods that we classify as savory. Indeed, it was Ikeda’s
(1908) observation that these foods have a distinct “savory” taste that led to his use
of the word umami to describe this.
The most concentrated levels of free glutamate are found in a wide variety of
foods, including some vegetables (e.g., mushrooms, potatoes, cabbage, corn, onion,
spinach), fruits (e.g., tomatoes, avocado), some fish and seafood (e.g., prawn, clam,
crab, oyster), seaweed, egg yolks, ripened cheeses, human breast milk, and fer-
mented soya beans and in lower concentrations in commonly consumed meats (e.g.,
beef, pork, chicken) (Kurihara, 2009; Ninomiya, 1998; Drake et al., 2007; Fuke &
Shimizu, 1993; Hajeb & Jinap, 2015; Ghirri & Bignetti, 2012). From a nutritional
standpoint, all of these provide valuable nutrients, but at first sight there appears to
5 Umami and Satiety 103
be no obvious strong nutritional link across these foods that could have driven the
evolution of umami as a preferred taste.
However, ribonucleotides are distributed rather differently. IMP in particular is a
constituent of most of the meat sources that provide most of the protein humans
consume (beef, lamb, pork, chicken, fish, and goat). Moreover, although the protein
found in those foods has a relatively weak taste profile in its natural state, once
hydrolyzed, the protein has a characteristic umami taste carried predominantly by
glutamate and ribonucleotides. Notably, the processes of aging, heating, and curing
these foods releases both free MSG and IMP, greatly enhancing the level of umami
taste (Sasaki et al., 2007; Rotola-Pukkila et al., 2015; Maga, 1994). This has led to
important speculation that our penchant for umami taste evolved at a time when
humans were switching to cooking as the primary process to prepare food for inges-
tion (Wrangham, 2017; Valente et al., 2018). Thus, in the form that we consume
these foods, umami may provide an oral signal of the likely presence of protein, an
idea first noted by Ikeda (1908).
It is therefore possible to develop a hypothesis that we evolved umami taste as a
way of detecting protein in cooked foods. However, unlike other basic tastes, umami
is strongly influenced by other flavor elements, making the sensing of umami more
complex than sensing MSG and ribonucleotides alone (Mouritsen et al., 2017). This
complexity of umami taste, with the strongest umami taste found only when sources
of MSG and IMP/GMP are combined with other flavor elements (McCabe & Rolls,
2007), may have had a particularly profound impact on the development of cuisine
worldwide, underlying the use of umami-rich seasonings (Mouritsen et al., 2017)
and shaping how we combine ingredients to better enable sensing of protein
(Ninomiya, 2015). The general idea that the overall savory character of a food is
then related to protein content has been supported by some studies that asked par-
ticipants to rate the savory character of foods and then explored how these ratings
related to actual protein content (van Dongen et al., 2012; Martin et al., 2014). Other
studies, however, suggest this relationship is relatively weak (Teo et al., 2018) or not
significant (Buckley et al., 2018). Overall, both the content of free glutamate and
umami-related nucleotides in foods and consumer evaluations of food products sug-
gest umami taste may function, at least in part, to locate protein in the diet, although
more work is needed to fully clarify this relationship.
5.2 Umami and the Regulation of Protein Intake
The argument developed in this chapter is that umami taste may go beyond a mech-
anism for simply sensing potential sources of protein in our diet to actually playing
a key role in the regulation of protein intake. To understand this, the evidence that
protein intake is regulated, followed by the effects of protein on satiety, is reviewed.
Then the literature that has examined how the presence of umami taste modifies our
experience of appetite is explored, focusing in particular on evidence that umami
may enhance satiety when protein is ingested. Studies that tested liking for umami
104 M. R. Yeomans
tastes as a function of protein need state provide further evidence for a role of umami
in satiety, suggesting that umami is liked more when we are in short-term need of
protein and, conversely, that liking is suppressed when we are protein replete.
5.2.1 Regulation of Protein Intake
Nutritionally, protein is one of the three main macronutrients and is an essential

component of the human diet, needed for the formation and repair of muscle tissue
and to provide essential amino acids to produce key neurotransmitters, hormones,
enzymes, and receptors, among other key cellular functions. Both fat and carbohy-
drate can be stored as reserves if intake exceeds current energetic needs, but protein
cannot be stored in the body. This inability may explain why the proportion of pro-
tein in the diet is remarkably similar in human cultures with very different dietary
habits. An analysis of the percentage of energy consumed as protein across 175
countries (Fig. 5.1: based on data published in Ritchie & Roser, 2017) gave an aver-
age of 11.1%, with more than 50% of countries having protein intake between
10.5% and 12.5% of total energy intake and only 10% of countries having more
than 12.6% of total energy from protein. Remarkably similar overall protein intake
was seen in countries where more than 65% of protein intake comes from animal
sources or 85% comes from plants. These data provide striking evidence of how
tightly controlled human protein intake is and adds credence to the idea that oral
sensing of protein in food, likely involving umami taste, is a critical part of appetite
control.
The evidence that protein intake is regulated more tightly than are fat and protein
has important implications for understanding our habitual food intake. According to
51
41
33
Frequency
21
16
5 4
2 1
0
0 - 6.5 6.5 - 7.5 - 8.5 - 9.5 - 10.5 - 11.5 - 12.5 - 13.5 - 14.5 -
7.5 8.5 9.5 10.5 11.5 12.5 13.5 14.5
Percentage of caloric intake from protein
Fig. 5.1 Average protein intake, as a percentage of total caloric intake, in 175 different countries
and dependencies (numbers at tops of columns). (Data from Ritchie and Roser (2017))
the protein leverage hypothesis, the absolute need for a certain level of protein in
our diet could drive overconsumption where available levels of protein are reduced
(Simpson & Raubenheimer, 2005; Gosby et al., 2014). A full evaluation of that
hypothesis is outside the scope of this chapter, but in the present context, the key
message is that protein intake is tightly regulated, and for that to be true, the body
has to have a mechanism for sensing and monitoring protein intake. Umami taste
offers a potential sensory component that could achieve this.
5.2.2 Protein: The Most Satiating Macronutrient?
Satiety could potentially be explained by a broader sensing mechanism that moni-

tors overall energy intake relative to energetic needs. At its simplest level, that prop-
osition suggests that energy is the key controlled variable, and the prediction then
follows that intake of the three main sources of energy (fat, carbohydrate, and pro-
tein) would have satiating effects, once matched for energy. However, humans gen-
erally experience greater satiety, that is, stronger suppression of appetite and
subsequent less ingestion at the next meal, after consuming protein-rich foods than
after energy-matched fat- or carbohydrate-rich foods (see Anderson & Moore, 2004;
Dhillon et al., 2016; Johnstone, 2013; Morell & Fiszman, 2017; Paddon-Jones et al.,
2008; Veldhorst et al., 2008 for reviews).
In this context, most short-term studies examining satiety use the preload design
(Yeomans, 2018), where participants ingest a fixed amount of different versions of
a test food, as the main manipulation, and their appetite is tested afterward to assess
satiety. A large number of preload studies using different foods and drinks, different
protein sources, and varying times between the preload and the ad libitum test meal
have shown greater satiety after protein than after other energy sources (e.g.,
Anderson & Moore, 2004; Bertenshaw et al., 2008; Chungchunlam et al., 2012;
Martens et al., 2013). Notably, however, a number of well-designed studies failed to
find any greater satiety for protein than for other macronutrients (e.g., Geliebter,
1979; Raben et al., 2003) and in some cases even found no evidence that any mac-
ronutrient source generated short-term satiety relative to a low-energy control (e.g.,
de Graaf et al., 1992). Although small methodological differences (study power,
satiety measures, etc.) could have influenced these outcomes, the variability in out-
come itself could suggest something important about how protein is sensed and
controlled. Crucially, to enable clear claims about protein, most protein-satiety
studies take great care to disguise the presence of protein at a sensory level. In doing
so, these studies may inadvertently remove the very signals the body uses to sense
and control protein intake—including, critically, umami taste.
This idea that the sensory characteristics of protein-rich products at least par-
tially explain the higher levels of satiety they generate was tested explicitly
(Bertenshaw et al., 2013). Their earlier studies had confirmed both that a whey-
protein-enhanced beverage was more satiating than an equicaloric carbohydrate-
enhanced beverage (Bertenshaw et al., 2008) and that the effects of whey protein on
106 M. R. Yeomans
satiety were dose dependent (Bertenshaw et al., 2009). However, the flavor charac-
teristics of whey protein (which includes an umami component) made it impossible
to fully disguise the addition of protein. Thus, they surmised that perhaps the greater
effectiveness of protein than carbohydrate to induce satiety may have been due to
the sensory characteristics cuing an expectation of protein, which in turn generated
the stronger satiety response. To test this, they contrasted the effects of the same
high-whey-protein preload with two alternative high-energy preloads: a carbohy-
drate preload adjusted to the same perceived thickness, with cream flavoring added
to match sensory characteristics of the whey preload, and a second protein preload
using a different form of whey protein that lacks these sensory characteristics
(Bertenshaw et al., 2013). Notably, when sensory matched, the high-sensory whey
and sensory-matched carbohydrate induced the same levels of satiety (evidenced by
lower-energy intake at a test meal; Fig. 5.2), but the equicaloric low-sensory protein
preload produced significantly less satiety. However, this study did not include any
specific manipulation of umami taste. Thus, studies that examine whether satiety is
greater in the context of umami taste can test the umami-satiety hypothesis.
5.3 A Role for Umami in Protein-Based Satiety?
How might the possible effects of umami on appetite be tested? To date, a number
of different approaches have been used, and although the picture is far from com-
plete, several lines of evidence suggest that umami may have some role in generat-
ing postmeal satiety. This research is described in the following sections, grouped
by the approach taken: human experimental preload-satiety studies with adult vol-
unteers, insights from the studies of feeding by human infants, and consideration of
how acute protein need state modifies responses to umami taste.
5.3.1 Umami-Enhanced Satiety: Human Experimental Studies
To date, 10 studies conducted with human adult volunteers, reporting 13 experi-

ments and representing research from 6 different research groups, have tested the
effects of added MSG or a combination of MSG with nucleotides including IMP on
satiety (see Table 5.1). These studies provide the most direct test of the idea that
umami enhances satiety and so warrant detailed evaluation. These studies are pre-
sented in chronological order since the findings changed as the design of studies
became more sophisticated.
The first study that directly assessed the effects of umami using a laboratory-
based satiety test in human volunteers looked at the effects of a relatively high
concentration of MSG (20% w/w) added to a minimal-energy beef consommé in
three related experiments (Rogers & Blundell, 1990). There was no evidence that
addition of MSG-enhanced satiety in these three experiments, with no effect on the
45
A) Control
40
35
Change Hunger Rating
30
25
20
15
10
5
0
-5
15 30 45 60 Time 75 90 105 120
B) Protein
45
40
35
30
25
20
15
10
5
0
-5
15 30 45 60 75 90 105 120
Time
C) Carbohydrate
45
40
35
30
25
20
15
10
5
0
-5
15 30 45 60 75 90 105 120
Time
Fig. 5.2 Recovery of hunger after consuming one of six versions of a naturally low-glutamate
soup (Masic & Yeomans, 2013): lower-energy (126 kcal) soup (a), higher-energy soup (310 kcal)
primarily with added protein (b), or a similar higher-energy soup (303 kcal) with added carbohy-
drate (c), either with (broken lines) or without (solid lines) added MSG
108 M. R. Yeomans
Table 5.1 Preload-satiety studies investigating effects of manipulated umami taste in human adult
volunteers
Umami
Study Experiment design manipulation Participantsa Key outcomes
Rogers and Effects of three 20% MSG 32 healthy No significant
Blundell preloads on rated volunteers difference in
(1990) appetite over 60 min: appetite or fullness
No preload or between MSG and
low-energy (<10 kcal) control soup
beef consommé with or
without nnn
Rogers and Effects of four preloads 10% or 20% 32 healthy No significant
Blundell on intake at a test meal MSG volunteers effects of added
(1990) consumed 30 min later: MSG on test meal
No preload or intake (trend for
low-energy (<10 kcal) small increase)
beef consommé with 0,
10, or 20% added MSG
Rogers and Effects of three 20% MSG 32 healthy No significant
Blundell preloads on intake at a volunteers effects of added
(1990) test meal consumed MSG on test meal
2 mins later, plus intake. More rapid
subsequent rated recovery of
appetite: No preload or appetite after the
low-energy (<10 kcal) soup with MSG
beef consommé alone
or with MSG
Luscombe- Effects of five preloads 0.6% MSG, 10 men, BMI No significant
Marsh on appetite, satiety- 0.25% IMP 26.5 ± 1.2, age effects of any MSG
et al. related hormones, and 44 ± 6 years; 12 manipulations on
(2009) intake at test buffet women, BMI rated appetite:
30 min later: Water 23.7 ± 1.4, age Significantly
control and four 32 ± 6 years greater food intake
isoenergetic test after the preload
preloads combining with MSG (but not
noodle soup + one or with MSG + IMP)
two filled rolls (overall than without either
nutrient content not additive
detailed: Portion sizes
adjusted to provide
20% of estimated daily
energy needs): Soup +
roll alone, + MSG, +
MSG/IMP, + MSG/
IMP sham-fed
(continued)
Table 5.1 (continued)

Umami
Carter et al. Effects of four 0.5% MSG, 35 healthy women: No effects of MSG
(2011) chicken-broth preloads 0.0013% BMI 21.6 ± 0.3, on test lunch
on appetite and test nucleotides age intake, which was
meal intake: Low- 24.7 ± 0.8 years reduced in the
energy (15 kcal) high-fat condition.
control, low energy + Significant
MSG, low energy + reduction in hunger
MSG and nucleotides, and desire to eat
or high energy after MSG at
(163 kcal; energy second preload
added as fat). Each
soup was consumed
twice, at 0900 and
1115 h, prior to test
meal at 1200 h
Finlayson Effects of equicaloric Not specified 30 healthy women: Lower intake of
et al. mushroom-flavored BMI = 22.7 ± 0.4, high-fat foods after
(2012) sweet, savory (+ MSG), age MSG-enhanced
or bland milk-based 21.9 ± 0.5 years savory than after
preloads on intake at sweet foods; no
buffet lunch 30 min other effects of
later savory
manipulation
Masic and Effects of six low-MSG 1% MSG 24 healthy men: Significantly
Yeomans vegetable soup BMI 24.0 ± 0.5, slower recovery of
(2013) preloads (covert age 21 ± 1.4 years hunger after MSG
nutritional than after control
manipulations) on rated in protein
appetite over condition; trend for
subsequent 2 h: Low slower recovery of
energy (126 kcal, hunger after MSG
control), higher-energy in low-energy
carbohydrate condition; no effect
(303 kcal), or of MSG in
higher-energy protein carbohydrate
(310 kcal), with or condition
without MSG
Masic and Effects of six low-MSG 1% MSG 35 healthy men: Reduced test meal
Yeomans vegetable soup BMI 22.0 ± 0.2, intake after protein
(2014a) preloads (covert age 21 ± 0.2 years but not after
nutritional carbohydrate
manipulations) on manipulation, with
intake at a test meal compensation for
45 min later: Low- covert energy
energy (126 kcal, enhanced by the
control), higher-energy presence of MSG
carbohydrate
(303 kcal), or
higher-energy protein
(310 kcal), with or
without MSG
(continued)
110 M. R. Yeomans

Umami
Masic and Effects of four 1% 11 men and 16 Significant
Yeomans low-MSG vegetable MSG + 0.25% women: BMI reduction in test
(2014b) soup preloads on intake IMP 22.7 ± 0.4, age lunch intake after
at a test meal 45 min 21.8 ± 0.6 years umami
later: Low-energy manipulation
(126 kcal, control) or relative to control
higher-energy protein and after protein
(310 kcal), with or relative to low
without MSG energy.
Significantly
higher
compensation for
added nutrients in
MSG (70%) than
for control (40%)
Imada et al. Effects of three 0.5% MSG, 86 healthy women: MSG, but not
(2014) low-energy (8 kcal) 0.0005% IMP BMI 22.2 ± 0.2, MSG + IMP,
chicken-broth preloads age significantly
on appetite and intake 37.9 ± 0.5 years reduced overall
at multi-item test meal energy intake at the
served 15 min later: test, with reduced
Broth alone (control), energy from sugar
broth + MSG, or broth and fat from the
+ MSG/IMP test buffet. No
significant effects
on rated appetite
Miyaki Effects of two 0.5% MSG 68 women, BMI Significantly lower
et al. low-energy (47 kcal) 28.6 ± 0.4, age food intake at test
(2016) soup preloads on test 38.8 ± 1.2 years lunch, but not at an
meal intake: Control afternoon snack,
soup or soup + MSG after soup + MSG
Anderson Effects of four 1.0% MSG 52 healthy young More sustained
et al. low-MSG vegetable men fullness after
(2018) soup preloads, plus protein + MSG
water control, on rated compared to all
appetite and test meal other conditions;
intake 120 min later: reduced desire to
Low-energy (281– eat in the
293 kcal, control) or low-energy MSG
higher-energy protein condition. No
(402–418 kcal), with or significant effects
without MSG of MSG on food
intake
(continued)

Umami
Anderson Effects of four 1.0% MSG 52 healthy young Lower subjective
et al. low-MSG vegetable men appetite after
(2018) soup preloads, plus protein + MSG
water control, on rated than in other
appetite, test meal conditions.
intake 120 min later, Reduced lunch
and selected satiety intake after protein
hormones: Low-energy + MSG but no after
(281–293 kcal, control) protein alone.
or higher-energy Increased release
protein (402–418 kcal), of insulin and
with or without MSG C-peptide and
reduced blood
glucose after
protein + MSG
Average participant age BMI are only reported here if detailed in the relevant paper
a
experience of appetite over the ensuing 2 h (experiment 1) and no significant effect

of 10% or 20% MSG on test meal intake (experiments 2 and 3). The only significant
effect on any of the measures of appetite was for more rapid recovery of hunger
after consuming the soup with 20% MSG followed 10 mins later by a meal (experi-
ment 3). Thus, these experiments provide no evidence of any effects of MSG on
satiety and, indeed, found instead limited evidence that MSG could stimulate appe-
tite, although the levels of MSG used were notably high and outside the physiologi-
cal range consumers would experience from normal foods.
The next study (Luscombe-Marsh et al., 2009) likewise failed to find any evi-
dence that umami caused satiety. Here, appetite and intake at a test meal were con-
trasted between a water control and a high-protein meal, with four variations:
unaltered, with added MSG (0.6%), with added MSG + IMP, or with the MSG/IMP
version sham-fed (i.e., chewed but not swallowed). There were no significant differ-
ences in changes in appetite ratings across time between the three high-protein
meals, all of which generated greater satiety (i.e., reduced rated appetite) than did
the water control or the sham-fed preload. Contrary to the predictions of enhanced
satiety, more was consumed after the meal with added MSG (but not added MSG +
IMP) than after the unaltered meal. Thus, the conclusion from these first two studies
would be that umami does not enhance satiety; rather, to the contrary, MSG might
cause a small increase in appetite. However, in the next study (Carter et al., 2011),
when a low-energy broth was consumed twice prior to a test meal, with umami
manipulated by the addition of MSG or MSG + IMP, MSG slowed the recovery of
hunger after the second soup preload. Although the umami manipulations had no
significant effects on food intake, this study did suggest a possible, albeit small,
effect of umami on satiety, and notably, this was seen at much lower levels of added
MSG than in the earlier studies. The approach in the next study was different:
Finlayson et al. (2012) tested whether consumption of a preload that was bland,
112 M. R. Yeomans
sweet, or savory in flavor modified food selection and intake at a subsequent buffet
meal. The relevance here was that MSG (level not reported) was used to modify the
flavor of the savory condition. Overall, there was little effect of the sensory manipu-
lations, but notably, intake of high-fat foods was significantly lower after the MSG-
enhanced savoury than the sweet preload.
Building on the findings of reduced recovery of hunger after an umami-enriched
soup (Carter et al., 2011), and the finding that carbohydrate could have the same
effects on satiety as protein if the sensory characteristics were similar (Bertenshaw
et al., 2013), Masic and Yeomans (2013) tested the effects of a realistic level of
MSG added to a low-energy, low-glutamate soup, and the same soup with either
added protein or added carbohydrate. As in an earlier study (Carter et al., 2011),
there was evidence that MSG could slow the recovery of hunger after ingestion, but
only after consuming the soup with added protein (Fig. 5.2), in contrast to the results
of Carter et al. (2011), who only manipulated MSG content of a low-energy (15 kcal)
broth. The implication is that the MSG signal appears to be most effective when it
is experienced in the context of actual protein ingestion. This might suggest that
umami acts as a signal of the likely presence of protein, thus aiding more efficient
processing of ingested protein, an idea further supported by a follow-up study that
examined test meal intake after low- and higher-energy soup preloads with and
without added MSG (Masic & Yeomans, 2014a). In that study, addition of protein
to a low-energy, low-glutamate soup reduced test meal intake more than did an
equicaloric carbohydrate preload. To further characterize satiety, the satiating effi-
ciency of the two macronutrient manipulations, with and without added MSG, was
calculated (Kissileff, 1984; Bellisle & Blundell, 2013). Satiating efficiency is calcu-
lated by determining what percentage of the energy difference between a treatment
and control preload is compensated for by reduced intake at subsequent meals.
Hypothetically, imagine someone consuming a low-energy (e.g., 100 kcal) preload
on 1 day and a high-energy (e.g., 400 kcal) on a second day. If they then consumed
300 kcal less at lunch after the high-energy than after the low-energy preload, the
satiating efficiency of the high-energy preload can be said to be 100%. In practice,
perfect compensation in preload-satiety studies is very rare (Almiron-Roig et al.,
2013; Chambers et al., 2015). Notably, when these values were calculated (Masic &
Yeomans, 2014a), compensation for the added energy was significantly greater after
the protein-rich soup with added MSG (62%) than in the equivalent carbohydrate
condition (24%). This study therefore added to the evidence that umami can enhance
satiety but that it does so particularly in the context of protein intake.
So far all of the umami-satiety studies in our laboratory had relied solely on the
manipulation of MSG content only. However, earlier it was noted that cuisine favors
combinations of ingredients that result in the presence of MSG + IMP, and the first
study to suggest there may be a role of umami in satiety included an MSG + IMP
manipulation (Carter et al., 2011). Therefore, Masic and Yeomans (2014b) exam-
ined the effects of MSG + IMP on protein-induced satiety by contrasting appetite
and intake after consuming a low-glutamate soup with added energy (principally as
whey protein) and a combination of MSG + IMP. In this case, there was evidence
both for the expected effect of protein on satiety, with less consumed after the
high-energy protein soup than after the low-energy soup, and an overall effect of
umami, with less consumed when umami was enhanced by the addition of MSG +
IMP in both the low-energy and protein-enriched preloads (Fig. 5.3). They again
calculated satiating efficiency and found that added protein plus MSG + IMP pro-
duced significantly greater compensation (70%) than did added protein alone (44%).
Since the series of studies in our laboratory, we are aware of a further three pub-
lications that have further investigated the possible effects of umami on satiety using
the preload-satiety model. In the first of these (Imada et al., 2014), participants
consumed a low-energy (8 kcal) chicken broth either alone (control), with added
A) Main Meal Intake

600
*
500
Calories Consumed (kcal)
*
400
300
200
100
0
Control Protein
B) Energy Compensation
100
90 *
Perecentage Compensation
80
70
60
50
40
30
20
10
0
No MSG/IMP Added MSG/IMP
Fig. 5.3 (a) Intake at lunch after consuming a low-energy (12 kcal) control soup (left) or a higher-
energy (310 kcal) soup, with energy mainly from protein (right), either with (white columns) or
without (gray columns) added MSG/IMP (Masic & Yeomans, 2014b). (b) Percentage of the pre-
load energy compensated for by reduced lunch intake after the high- relative to the low-protein
preload in the control and MSG/IMP conditions. * p < 0.05
114 M. R. Yeomans
MSG (0.5%), or with added MSG + IMP prior to a multi-item buffet meal. Overall
energy intake at the meal was lower after consuming the soup with added MSG, but
not with added MSG + IMP, than in the control condition, due primarily to the
reduced selection and intake of sweet and high-fat snacks. Thus, this study did sug-
gest some direct satiating effects of a low concentration of MSG but found this in
the context of a minimal-energy broth. A subsequent study by the same group tried
to extend the findings from the study by Carter et al. (2011) using the two-soup
intervention model in overweight individuals, with a soup naturally low in gluta-
mate (Miyaki et al., 2016). Again, addition of 0.5% MSG to this soup reduced sub-
sequent food intake at a test meal and also reduced rated hunger between soup
ingestion and the start of the meal.
In the most recent two experiments to examine the effects of umami on satiety,
Anderson et al. (2018) attempted to replicate and extend the earlier findings of
Masic and Yeomans (2014a). The key differences were the use of a slightly larger
preload (500 mL instead of 300 mL), a longer gap between preload and test meal
(120 min instead of 45), and measurement of satiety-related hormones as additional
measures of satiety. In their first experiment, the high-protein soup with MSG sus-
tained fullness for a longer period than did all other treatments, but MSG had no
significant effects on intake at the test meal. In their second experiment, subjective
appetite was significantly lower after the protein soup with added MSG than after
all other conditions, and intake at the next meal was significantly lower for the pro-
tein + MSG but not protein-alone conditions than in the lower-energy and water
control conditions. Both experiments thus found some evidence to support a role for
umami in satiety, replicating earlier studies. The second experiment also found
some changes in physiological markers that further supported these behavioral sati-
ety findings: the protein + MSG preload but not the protein-alone preload resulted
in increased levels of insulin and C-peptide (an early-stage marker for insulin syn-
thesis) and also lowered blood glucose.
At face value, of the 13 experiments examining effects of MSG on satiety, 9
reported at least some evidence supporting a role for umami in satiety, and of the 4
that did not, 3 were from the first study to examine this question. Closer inspection
of the designs of these studies gives clues to why some did not find evidence of
umami-enhanced satiety. The first key design issue appears to be the dose of MSG
used to manipulate umami: the dose used in the earliest three experiments (Rogers
& Blundell, 1990) was considerably higher (10–20%) than in the more recent
experiments finding evidence of satiety (typically 0.5–1.0%). The levels of MSG in
those early studies were considerably higher than those found in normal food prod-
ucts, and it is notable that studies using more realistic levels of MSG found evidence
of satiety enhancement. This further implies that satiety is not enhanced as a linear
function of MSG concentration. The second design issue may be the size of the
preload: all of the successful studies used relatively low-energy preloads (maximum
of ~300 kcal), nearly always as a soup manipulation. In contrast, the study by
Luscombe-Marsh et al. (2009), which found no significant effect, used a much
larger preload, comprising soup plus one or two savory rolls, with the portion size
adjusted to deliver 20% of the estimated energy requirements for each participant.
The effects of umami may have been masked by the much stronger satiety from the
larger preload. This in itself raises important questions about the nature of the sati-
ety signal umami generates, discussed in depth further below. A final issue worth
noting is that many studies did not fully control the level of umami taste in the con-
trol conditions of these studies, which hinders interpretation. For example, if the
level of umami in the control was already sufficient to signal the possible presence
of protein, then additional MSG might not add any new information that could
affect appetite.
5.3.2 Glutamate and Satiety in Human Infants
Although it is less possible to conduct direct studies of the role of umami in satiety
in infants and children, further insights into the potential impacts of umami on sati-
ety have been provided by studies exploring differences in weight gain associated
with breast-feeding and bottle-feeding practices in humans. In relation to umami
taste, one of the most surprising observations is that human breast milk has as much
as 19 times the free glutamate content of cow’s milk (Van Sadelhoff et al., 2018).
That observation allowed researchers to consider what role umami might have in
control of infant feeding (Mennella et al., 2011; Ventura et al., 2012, 2015), in the
context of the well-established finding that human infants who are bottle-fed a diet
of standard cow milk formula (CMF) typically gain more weight than do babies
who are breast-fed (reviewed by Appleton et al., 2018).
In addition to the difference in free glutamate, there is a notable difference in
overall protein content of the two main milk sources fed to human infants: CMF has
more protein overall than does human breast milk, and consequently, formula-fed
infants have much higher protein intake (55–80% more, adjusted for body weight;
Alexy et al., 1999) than do breast-fed infants. Because protein is more satiating than
other macronutrients, we might expect that CMF would be more satiating than is
breast milk and that the faster weight gain seen in CMF-fed infants cannot be attrib-
uted to a lack of satiety from feeding. However, while CMF has higher overall
protein content, it has lower overall levels of free amino acids, most notably gluta-
mate. When the growth rate of human infants was contrasted between standard
CMF and a formula with protein that had been extensively hydrolyzed (EHF),
which consequently had a much higher free amino acid content (Mennella et al.,
2011), those fed on EHF had more normative weight gain than those fed standard
CMF, and their growth rates were in line with predictions for breast-fed infants (for
further details, see Chap. 5 in this volume). Hydrolyzing the protein in the formula
had a particularly large effect on levels of free glutamate, with 106.5 mg/100 mL in
the hydrolyzed formula compared to just 1.8 mg/100 mL in CMF. These short-term
effects were subsequently verified in a larger randomized controlled trial to more
fully determine the direct impact of standard CMF and EHF on infant growth
(Mennella et al., 2018). Replicating the initial finding, that study found evidence of
sustained slower growth in body weight for infants fed with EHF, which persisted
116 M. R. Yeomans
across the first year of life, because infants ingested more CMF than EHF, driven at
least partly by a reduced intake of EHF, which was more satiating.
More remarkably, the same group extended this work to contrast whether the
presence of free amino acids affected actual infant feeding (Ventura et al., 2012).
Infants attended the laboratory on three occasions to consume two meals with each
of three formula diets: CMF, EHF, and CMF with added MSG. Infants fed both with
EHF and MSG-enhanced formula consumed less at the first meal than they did with
CMF, and a longer interval elapsed before they signaled readiness for their second
meal, and they did not compensate by consuming more at that second meal. Thus,
both reduced immediate intake and delayed demand for the second meal are consis-
tent with an effect of umami taste on satiety in human infants, adding further weight
to the idea that umami has a clear role in regulating human appetite. A subsequent
study explored the feeding style of infants fed standard and CMF with added gluta-
mate (Ventura et al., 2015). Infants consumed less of the glutamate-enhanced CMF
and tended to feed for shorter times. Although detailed analysis of feeding behavior
(from video analysis) found few differences between diets, there was evidence that
the distinct end-of-meal behaviors seen later in meals became evident sooner with
the glutamate-enhanced formula. The reduction in intake, shorter feeding duration,
and earlier switch to end-of-meal behaviors might be consistent with increased sati-
ety from the glutamate-enhanced CMF.
5.3.3 Protein Need State, Satiety, and Liking for Umami Taste
The evidence to date suggests a likely role for umami taste in satiety in humans but
is far from conclusive. The inconsistent results among studies that explicitly
explored umami taste and satiety in particular (see Table 5.1) suggest that any such
role of umami may be specific to very defined conditions, and so whether this real-
istically contributes to satiety in real life may be questionable. However, if umami
acts as a signal for the likely presence of protein, then biologically it would make
sense for sensitivity to that umami signal to also vary depending on individual pro-
tein need state. If so, the variability in study outcomes could relate to differences in
acute and habitual protein intake across study designs and individual participants.
As discussed earlier, regulation of protein intake appears much more tightly con-
trolled than fat and carbohydrate, and this may in turn mean that sensitivity to
umami taste, including its effects on satiety, may also vary with both habitual pro-
tein intake and acute protein need. Thus, the final set of studies explored in this
chapter tested how responses to umami taste and/or wider savory evaluations vary
in relation to both acute and longer-term protein intake. The focus again is on human
studies because of the suggested evolution of a specific role for umami coinciding
with when humans first started to cook food (Hartley et al., 2019; Valente et al., 2018).
In support of responses to MSG varying with protein need state, a number of
older studies in humans reported stronger preferences for foods with higher MSG
content in both adults (Murphy, 1987) and children (Vazquez et al., 1982) when in
protein-deficient states or with poor nutritional status compared to well-nourished

controls. Some studies have also reported that liking for savory flavors varied with
acute protein status. In this context, one study in particular suggested that partici-
pants’ ability to respond to experimentally manipulated acute protein deficit (by
provision of low-protein breakfasts) has to be learned: participants who were acutely
protein deprived came to prefer a dessert flavor that was previously paired with the
delivery of more protein relative to participants tested and trained in the absence of
acute protein lack (Gibson et al., 1995). That study did not specifically manipulate
umami taste, but the demonstration that acute protein need heightened sensitivity to
sensory characteristics of a protein-rich product fits the wider idea that umami pro-
motes protein choice in the protein-deprived state. Specific evidence that acute pro-
tein deprivation modifies the response to MSG came from a study that examined
responses to umami, salty, and sweet tastes in participants in relation to acute pro-
tein need (by manipulating earlier protein intake) and habitual protein intake (Masic
& Yeomans, 2017). In that study, acute protein deprivation increased liking regard-
less of MSG concentration (Fig. 5.4a), and crucially, this effect depended on habit-
ual protein intake (Fig. 5.4b). Here, liking for the strongest (1%) MSG soup
decreased as a function of habitual protein intake after consuming a protein-rich
breakfast, but not after the low-protein or baseline (habitual) breakfast. Because the
high-protein breakfast would have been predicted to generate higher satiety, the
implication is that expression of liking for umami is suppressed by protein-induced
satiety. This suggests that umami’s role in appetite control is fine-tuned to control
protein intake, with increased preference for umami driving protein intake in the
context of either habitual low-protein intake or acute protein need. Moreover, if the
Fig. 5.4 How acute and habitual protein intake modified hedonic responses to umami taste (Masic
& Yeomans, 2017). (a) Overall changes in liking from baseline for a low-glutamate soup with 0%
(white columns), 0.6% (gray columns), or 1.0% (dotted columns) added MSG evaluated 2 h after
consumption of a low- or high-protein breakfast. * p < 0.05. (b) Absolute liking for soup with 1%
added MSG as a function of habitual protein intake on the baseline (normal breakfast) day (black
line) and after days when participants consumed a specific low-protein (dark gray line) or high-
protein (light gray line) breakfast
118 M. R. Yeomans
Fig. 5.4 (continued)
body monitors protein intake, at least in part, through experience of umami taste,
then increased exposure to umami taste may in turn result in reduced preference for
umami, an idea supported by a recent study where repeated exposure to umami
resulted in a generalized decline in liking for umami taste (Noel et al., 2018).
5.4 Possible Models for Umami-Enhanced Satiety

in Humans
If umami taste does have a role in short-term regulation of food intake, particularly
associated with satiety in the context of regulation of protein intake, a key question
is how this might work. The original idea that umami taste evolved to allow us to
sense protein in our diet (Ikeda, 1908) would not necessarily suggest that umami
should play a role in satiety: umami might direct food choice to protein-rich sources,
but the control of intake could be achieved through other satiety processes. However,
as discussed in detail above, intake of protein appears to be much more tightly regu-
lated than intake of other macronutrients. Thus, the effects of umami on satiety
might suggest umami taste has a role beyond food choice to help regulate protein
intake. How might that regulation be achieved?
Protein has been shown to be more satiating than other macronutrients, but this
is most evident where the sensory characteristics of the ingested food include cues
that predict the likely presence of protein (Bertenshaw et al., 2013). Umami taste
may be one of the most reliable predictors of protein in the context of the cooked
foods that most humans consume. This is supported by the positive findings of most
studies that examined the effects of umami taste on satiety and included a protein-
rich manipulation (Table 5.1). Although these effects were relatively subtle, these
studies found at least some evidence (in terms of altered intake and/or enhanced
subjective satiety) that the combination of MSG + protein was more satiating than
would be predicted from the sum of the separate effects of protein and MSG alone.
Although how umami taste magnifies the satiating effects of protein is not clear, two
possible models are presented here that might be explored in future research.
The first model builds on increasing evidence that the orosensory experience of
foods generates consumer expectations about the level of satiety they will experi-
ence postingestion (Chambers et al., 2015). There is now a wide variety of evidence
that both sensory and cognitive characteristics of ingested products can modify
these satiety expectations and that these expectations then interact with actual nutri-
ent consumption to generate satiety. Beyond umami, at a sensory level, two exam-
ples are (1) where perceived thickness and creaminess of a beverage generated
predictable expectations about satiety (McCrickerd et al., 2012), and these same
expectations then modified the degree of satiety generated by a disguised nutrient
load (Yeomans & Chambers, 2011), and (2) where differences in oil-droplet size
likewise altered satiety expectations (Lett et al., 2015) and actual satiety (Lett et al.,
2016). To date, studies have not explicitly tested how manipulation of umami modi-
fies satiety expectations, but the prediction would be that the increasing savory char-
acteristics of umami-rich foods would generate stronger satiety expectations.
However, if umami simply generated satiety expectations, then any nutrient-rich
food that has some level of umami taste should be more satiating than the same
nutrients without umami taste. What is more complicated with umami is that the
effects seem to be protein specific, which argues against a generalized expected
satiety model. However, if the expectations generated by umami were confirmed by
protein sensing in the gut postingestion, this model offers a plausible account. It is
now well established that the TAS1R1 and TAS1R3 receptor units that underlie
umami taste when glutamate is detected in the mouth are also present throughout
the gastrointestinal tract (Kondoh et al., 2009; San Gabriel & Uneyama, 2013),
although as with other postoral taste receptors, activation of these receptors does not
result in our “tasting” food in our gut. Instead, it is widely believed that these gut-
based receptors monitor nutrient levels (e.g., Shirazi-Beechey et al., 2014; Raka
et al., 2019), perhaps fine-tuning digestive processes depending on the balance of
ingested nutrients.
If the impact of umami on satiety was mediated through expectations, then we
would predict that umami taste would enhance behavioral measures of satiety
expectations and have measurable effects in brain areas associated with these expec-
tations. To date, there has been no detailed evaluation of effects of umami on
expected satiety, and likewise, the neural representation of expected satiety has not
been elucidated. However, a number of studies with human volunteers have looked
at neural responses to umami taste, which allow some inference of the plausibility
that umami enhances protein-induced satiety by generating expected satiety. Early
studies established that umami taste is represented in the primary and secondary
gustatory cortex, incorporating the anterior insula, frontal operculum, and the orbi-
tofrontal cortex (OFC: e.g., Schoenfeld et al., 2004; McCabe & Rolls, 2007; Singh
et al., 2015). The OFC has also been shown to have a role in sensory-mediated
satiety (O’Doherty et al., 2000). More importantly, the OFC has been shown to have
120 M. R. Yeomans
a critical role in predicting reward value (e.g., O’Doherty, 2007; Hare et al., 2008;
McDannald et al., 2012), and because satiety expectations are neural predictions of
the potential impact of ingestion on satiety, it would be predicted (but as yet not
tested) that OFC neurons would play a critical role in satiety expectations. Thus,
although no studies to date have formally tested an expected satiety model of
umami-enhanced satiety, current knowledge of encoding of umami taste and broader
prediction are both consistent with an expectation-based model.
An alternative to the expectation-mediated effects of umami on satiety would be
a more direct effect of umami on the physiological systems known to be involved in
satiety. Of principal interest are the specific hormones released in the gut in response
to nutrient ingestion (see Chaudhri et al., 2008; for reviews Zanchi et al., 2017),
which include cholecystokinin (CCK), pancreatic polypeptide, glucagon-like pep-
tide, and polypeptide-YY. CCK is of particular interest, since studies in vivo and in
nonhuman animals have provided some evidence that stimulation of the intestinal
TAS1R1 + TAS1R3 complex can stimulate CCK release (Daly et al., 2013; Tian
et al., 2019). If this is the cause of the enhanced satiety seen in some human appetite
studies, then it would be predicted that circulating CCK levels would be raised after
consuming umami-tasting foods, particularly in the context of protein consumption.
In this context, it is notable that one study in human volunteers found evidence of
reduced hunger, but not increased CCK, following intragastric infusion of umami
tastants (Van Avesaat et al., 2015). A more recent study examined effects of umami
taste and ingestion on glucagon-like peptide and again found no evidence of any
change in hormone levels (Anderson et al., 2018). Thus, the only two studies that
examined changes in satiety hormones after umami did not find evidence in humans
to support the suggested effects of umami on CCK release reported elsewhere.
Overall, while a direct effect of umami taste on gut-based satiety signaling cannot
be discounted, to date there is no evidence for this.
Finally, although the expectation-based and gut signaling-based models of
umami-based satiety are discussed here as separate potential models, in practice
control of appetite is multifaceted and integrated. Notably, there is increasing evi-
dence of top-down control of gut-based satiety signaling (e.g., Crooks et al., 2021)
and, in particular, that sensory characteristics of products that generate satiety
expectations may also enhance gut-based release of CCK (Yeomans et al., 2016).
5.5 Uncertainties and Future Directions
This chapter reviews evidence that umami may enhance satiety in humans and iden-
tifies important shortcomings in our current knowledge. The wider idea that umami
is an alimentary taste (Keast et al., 2021) that may have evolved in humans in the
context of our use of fire to modify protein prior to ingestion complicates the inves-
tigation of umami-induced satiety since it implies that the role of umami in humans
may differ from that in other species. Usually findings in humans are verified by
more detailed physiologically based studies in nonhuman animals, but in this case
such studies may not be appropriate tests of the role of umami in satiety in humans.
Instead, more detailed human studies are needed to meet specific shortcomings in
the literature, including (but not limited to) the effects of umami taste on satiety
expectations, more detailed evaluations of neural responses to umami taste in rela-
tion to satiety, and more highly powered and sustained studies of the effects of
umami-enriched products on satiety. A further complication is that many products
already have appropriate levels of satiety signals that match the relevant nutrient
content; adding umami taste in that context could lead to a mismatch between the
sensed taste and nutrient content, an issue that has been noted in the context of
sweet taste and carbohydrate-based calories (Veldhuizen et al., 2017).
5.6 Conclusions
This chapter evaluates the evidence that umami taste may impact satiety in humans.
The evidence that umami predicts protein, as originally suggested by Ikeda (1908),
does not look convincing until the cooked state of the food humans ingest is taken
into consideration. In the context of cooked food, the idea that umami predicts pro-
tein is plausible and also fits with evidence of how the system of sensing umami
taste evolved. Although not all studies reviewed found evidence of enhanced satiety,
the balance of evidence, particularly where the level of umami enhancement is
physiologically realistic, does suggest a subtle effect of umami on satiety. In par-
ticular, the evidence suggests that umami increases the degree to which protein is
satiating, which may explain why protein has been found in general to be more
satiating than other macronutrients. Future research is needed, however, to under-
stand the mechanisms through which umami enhances satiety. This chapter offers
the suggestion to focus on two models, one based on prediction error and expecta-
tions and the other on physiological satiety cues.
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Martin R. Yeomans is Professor of Experimental Psychology at the School of Psychology,

University of Sussex, in Brighton.
His research centers on the psychology and physiology of motivational controls of eating and
drinking, focused originally on basic physiological controls but more recently concentrating on the
role of learning. Current work is focused in particular on an integrated approach to understanding
eating motivation, combining cognitive, sensory, and nutritional signals. These factors in turn
impact on food choice and on causes of overeating. He has published over 150 journal articles and
contributions to specialist books.
Chapter 6
Umami Taste: Inborn and Experiential
Effects on Taste Acceptance and Satiation
During Infancy
Ana San Gabriel and Julie A. Mennella
The sensation of taste, which has been classically delineated into the five basic taste
qualities of sweet, sour, salt, bitter, and umami (Beauchamp, 2019), has attracted
great interest in recent years as a major determinant of what we eat—an overlooked
aspect of nutrition (Kershaw & Mattes, 2018). Taste plays a critical role as the gate-
keeper of what enters the body, guarding against ingestion of dangerous substances
(e.g., bitter-tasting poisons) (Glendinning, 1994) while encouraging consumption
of foods important for growth and development (Kurihara, 2015; Gabriel et al.,
2018), including mother’s milk, readily available glucose in energy-containing
plants (e.g., sweet fruits) (Beauchamp, 2016), a needed mineral (salt) (Beauchamp,
1987), and proteins (umami)—particularly the taste of the amino acid L-glutamate
(Glu) and 5′-ribonucleotides (Beauchamp, 2009). In the context of feeding, these
taste sensations naturally co-occur with other sensory modalities, including olfac-
tion and chemesthesis.
This chapter focuses on umami taste in infancy. As a basic taste, the scientific
investigation on umami has received perhaps the least attention in developmental
studies, especially when compared to the ontogeny of sweet and salty tastes
(Mennella et al., 2016a; Beauchamp & Mennella, 2009). We provide an overview of
the basic biology of umami taste, from the distribution of umami taste receptors
throughout the oral cavity and gastrointestinal system to its role in flavor, palatabil-
ity, and food intake. We then summarize the scientific evidence on early routes of
umami exposure and children’s inborn and learned responses to its taste and satiat-
ing properties. We focus on the first year, when infants make the drastic transition
from eating an all-liquid diet of human milk, artificial milk (infant formula), or
A. S. Gabriel
Global Communications, Ajinomoto Co., Inc, Tokyo, Japan
J. A. Mennella (*)
Monell Chemical Senses Center, Philadelphia, PA, USA

https://doi.org/10.1007/978-3-031-32692-9_6
128 A. S. Gabriel and J. A. Mennella
both, to one containing both liquid and solid foods (Grummer-Strawn et al., 2008).
This body of scientific evidence derives from a variety of precise and detailed mea-
surements of infants’ orofacial responses and well-controlled paradigms that mea-
sured their behavioral responses, intake, and satiation (Ventura & Mennella, 2017).
Often the umami stimulus studied was L-glutamate, in the form of the sodium salt
monosodium glutamate (MSG).
6.1 Umami Taste in the Oral Cavity and Alimentary Canal
One century after Kikunae Ikeda’s description of the fifth basic taste of umami
(Ikeda, 2002), scientific breakthroughs revealed its perception is mediated by acti-
vation of heterodimeric G-protein-coupled receptors (GPCRs) encoded by mem-
bers of the type 1 family of taste receptor genes, TAS1R1 and TAS1R3 (Nelson et al.,
2002; Bachmanov et al., 2016) (see Chaps. 1 and 2). In addition to the TAS1R
genes, other GPCRs act as umami receptors: truncated taste versions of the metabo-
tropic glutamate receptors (mGluRs) found in many neuronal cells, mGluR4
(Chaudhari et al., 2000a) and mGluR1 (Chaudhari et al., 2000a; Yasumatsu et al.,
2015; San Gabriel et al., 2009). These umami receptors, located throughout the oral
cavity and alimentary canal, are activated only by free amino acids (FAAs), not by
amino acids bound in the form of proteins (Keast et al., 2021). The locations of
these receptors underlie their different roles in signaling the presence of taste-
reactive FAAs, which not only generate the perception of umami taste but also mod-
ulate complex digestive processes such as gastric emptying and satiety.
6.1.1 Oral Cavity and Taste Psychophysics
The T1R1 + T1R3 heterodimer receptor and mGluRs are located at the tip of taste
cells distributed across the tongue (Yasumatsu et al., 2015). When activated by free
L-amino acids (e.g., free Glu), 5′-ribonucleotides, or salts of glutamic acid (e.g.,
MSG), these receptors transmit signals to the brain. The taste of Glu is synergisti-
cally enhanced by the sodium salts of 5′-inosine monophosphate or 5′-guanosine
monophosphate.
Psychophysical studies revealed that MSG imparts a “savory” and “satisfying”
sensation (Rogers & Blundell, 1990), increasing the palatability of a variety of
foods, due in part to its ability to block bitter tastes (Mennella et al., 2014a).
Beauchamp (2009, 2019) suggests that umami also increases palatability by func-
tioning as a flavor enhancer (Beauchamp, 2009; Hartley et al., 2019), but the food
matrix in which umami-tasting compounds are experienced is important. Although
unpalatable when tasted alone or in aqueous solution, umami compounds are palat-
able in broth and improve the flavor of a variety of foods and flavors (Beauchamp,
2009; Yeomans et al., 2008; Yamaguchi & Takahashi, 1984), imparting a pleasant
6 Umami Taste: Inborn and Experiential Effects on Taste Acceptance and Satiation… 129
“mouthfeel” sensation of fullness (Prescott, 2004). Glu occurs naturally in many

foods, such as meats, cheeses, broths, and tomatoes, and imparts a savory taste
(Gabriel et al., 2018; Ninomiya, 2003). It is notable that many cultural food tradi-
tions intended to preserve food or increase its nutritional value (e.g., extraction,
curing, aging, fermentation) often increase the food’s FAA, including Glu, content
(Ninomiya, 2015).
6.1.2 Alimentary Canal and Feeding Behaviors
Outside of the oral cavity, T1R1 + T1R3 receptors and mGluRs are scattered widely
along the lower alimentary canal, comprising the stomach (San Gabriel et al., 2007)
and small intestine, where they regulate digestion, absorption, and metabolism of
nutrients, especially amino acids (Keast et al., 2021). Although perception of umami
taste in the mouth is a conscious sensation indicating the presence of free Glu in
foods, the detection of Glu in ingested food by the gastrointestinal tract is not
(Hartley et al., 2019). Instead, when umami compounds bind to receptors in the
gastrointestinal system, nerves are activated, hormones are released, or both
(Kondoh et al., 2009; Daly et al., 2013; Shirazi-Beechey et al., 2014).
Electrophysiological studies have revealed that binding of MSG to the stomach lin-
ing, probably via stomach mGluR1, can activate the vagus nerve by releasing bioac-
tive molecules such as serotonin (Uneyama et al., 2006), whereas molecular and
physiological studies suggest that free Glu in ingested food potentiates gastric
digestion and emptying of amino acid-rich foods (San Gabriel et al., 2007; Zolotarev
et al., 2009). In the intestine, the binding of free Glu to T1R1 and T1R3 receptors
appears to cause secretion of the hormones ghrelin and cholecystokinin, which are
known to impact gastric emptying and/or satiation (Keast et al., 2021; Vancleef
et al., 2015).
Brief feeding studies in healthy adults have shown that, when added to foods
such as broths and vegetables, MSG imparts a “savory” and “satisfying” sensation
that enhances palatability (Rogers & Blundell, 1990; McCabe & Rolls, 2007; Rolls,
2009; Carter et al., 2011). However, results on its effect on satiation (i.e., energy
intake within a meal) and satiety (i.e., energy intake during a subsequent meal) in
adults have not being consistent, perhaps due in part to the wide range of experi-
mental paradigms and participant ages and health status (Keast et al., 2021) (see
Chap. 4). While some studies on healthy, noninstitutionalized adults reported that,
when added to a food, MSG increased satiation and reduced intake within a meal or
increased sensations of fullness (Imada et al., 2014; Miyaki et al., 2016), others
reported no effects (Rogers & Blundell, 1990; Luscombe-Marsh et al., 2009;
Anderson et al., 2018).
6.2 Inborn Responses to Umami Taste
At birth, human infants are well equipped to convey a range of hedonic responses to
tastes, odors, and complex flavors. As reviewed by Forestell and Mennella (2017),
facial expressions in particular play an important adaptive role, allowing dependent
infants to convey information to caretakers about the sensations they are experienc-
ing in their oral cavity. Head turning away and displays of gaping and puckering in
response to bitter and sour tastes are often recognized as signals of disgust or rejec-
tion, whereas orientation toward the food, faster sucking, and smiling are often
interpreted as a signals of liking, encouraging feeding of that particular food.
In the 1970s and 1980s, a series of pioneering studies by Steiner, Ganchrow, and
colleagues revealed that, within hours after birth, newborns respond with character-
istic, differential orofacial responses when small quantities of sweet-, sour-, bitter-,
or umami-tasting liquids were placed on their tongue (Steiner, 1979, 1987;
Ganchrow et al., 1983). Similar to infants’ reactions to the taste of sugars, the facial
responses to the taste of umami appear to be primarily inborn and preferred (Steiner,
1987). Newborns responded with increased sucking and mouth movements and
relaxation of their face when tasting soup broth containing 0.1% and 0.5% MSG but
rejected MSG when presented in water solutions. Research on older infants
(2–24 months) confirmed the rejection of Glu in plain aqueous solutions (Beauchamp
& Pearson, 1991) and the palatability of umami in the context of a food. Both well-
nourished and malnourished 2- to 24-month-old infants preferentially ingested soup
broth containing MSG relative to soup broth alone (Beauchamp & Pearson, 1991;
Beauchamp et al., 1987).
This body of research highlights important aspects of the taste stimuli and meth-
odological approaches needed to examine umami taste palatability. First, it is diffi-
cult to interpret findings when children are presented with only umami taste in plain
aqueous solution (Schwartz et al., 2009). Rather, umami must be experienced in the
context of other chemosensory stimuli when testing individual differences in the
hedonics of umami taste in pediatric and adult populations (Beauchamp, 2009;
Forestell & Mennella, 2017; Steiner, 1987). As suggested by Beauchamp (2009,
2019), umami functions as a flavor enhancer, increasing the palatability of flavors it
is mixed with (Hartley et al., 2019). Second, when phenotyping orofacial taste reac-
tivity, infants should not see the facial expression of their mothers during testing
since they are sensitive to and mimic these facial expressions (Gunnar & Stone,
1984). Third, measurements of intake and acceptance should be based on infants’
behavioral signaling of hunger and satiation (i.e., infant-led feeding paradigms)
(Forestell & Mennella, 2017; Ventura et al., 2015) and not determined by mothers
or experimenters, because they may under- or overfeed by not attending to the
infant’s satiation cues (Crow et al., 1980; Li et al., 2010).
6.3 Early Experiences with Umami: First Foods
Like the prenatal diet (amniotic fluid), the early postnatal diet is unique in that it is
typically solely liquid based, consisting of human milk, infant formula, or both. The
protein in these first foods supports infant growth, immune function, and behavioral
development by supplying nitrogen and essential and semi-essential amino acids
(Dewey et al., 1996). These first foods—amniotic fluid, human milk, and infant
formula—as well as a variety of foods, from vegetables to meats, naturally contain
FAAs, including free Glu (Yamaguchi & Ninomiya, 2000).
Infants’ experiences with the taste of FAAs, and Glu in particular, can vary con-
siderably. Table 6.1 summarizes the variation in the total FAA and free Glu content
in amniotic fluid, human milk at different stages of lactation, and infant formulas
from a range of studies. Although measures of FAAs in human fluids date back to
the 1940s (Beach et al., 1941), we included only studies that used chromatography
and spectrophotometry measures, omitting earlier ones that used microbiological
Table 6.1 Total free amino acids (FAAs) and free glutamate (Glu) content of amniotic fluid,
human milk, and infant formulaa
Concentration (μmol/L)
Total FAAs
Sample (number of FAA)b Free Glu Selected References
Amniotic fluid
First and second trimesters 2036 (17) 149 Cockburn et al. (1970)
1803 (16) 261 Lopez Ramon y Cajal et al.
(2007)
2230 (16) 149 Reid et al. (1971)
Third trimester 1070 (16) 48 Reid et al. (1971)
1573 (16) 107 Levy and Montag (1969)
Human milk
Colostrum 3321 (19) 1090 Zhang et al. (2013)
Transitional milk 2416 (19) 960 Zhang et al. (2013)
Mature milk 2481–2941 (19) 1175– Zhang et al. (2013)
1529
Infant formula
Cow milk 523–864 (20) 86–109 Ventura et al. (2012a)
Soy 19,33–2450 (20) 0–11 Ventura et al. (2012a)
Partial protein hydrolysate 2329 (20) 113 Ventura et al. (2012a)
Extensive protein 80,375–85,445 (20) 7472– Ventura et al. (2012a)
hydrolysate 8226
a
Values represent normal pregnancies for amniotic fluid and term births for human milk. Data are
presented as ranges of means when from more than one publication, from two stages of lactation,
or from more than one brand of a given type of infant formula
b
Total concentration based on the sum of all FAAs; numbers in parentheses indicate number of
different FAAs summed
methods. Note that different studies used different numbers of FAAs to calculate
total FAAs, ranging from 16 to 20 individual FAAs.
6.3.1 Amniotic Fluid
Much of the early research on amniotic fluid content aimed to help diagnose certain
genetic disorders associated with aminoaciduria (Emery et al., 1970) or to identify
biomarkers of inborn errors of metabolism (Reid et al., 1971) or placental dysfunc-
tion (Lopez Ramon et al., 2007). During normal pregnancies, FAA levels change
dynamically in amniotic fluid (Reid et al., 1971) (see Table 6.1). In general, FAA
concentrations in amniotic fluid and maternal plasma are higher during early preg-
nancy and lower in the third trimester and near term (Emery et al., 1970; Reid et al.,
1971; Lopez Ramon et al., 2007; Athanasiadis et al., 2011; Levy & Montag, 1969;
Cockburn et al., 1970). Such changes in part reflect the exchange of maternofetal
fluids as the placenta and the fetus develop (Reid et al., 1971; Lopez Ramon et al.,
2007). Amino acids are actively transported across the human placenta, mediated by
specific transporters in plasma membranes (Jansson, 2001), and are also synthe-
sized and metabolized through the placenta (Jansson, 2001; Cetin, 2001).
The time course and trajectory of changes in amniotic fluid during pregnancy
differ among the individual FAAs (Reid et al., 1971; Guadalix et al., 1975). Overall,
alanine was the most abundant amino acid in the amniotic fluid during gestation
(Lopez Ramon et al., 2007). Although less abundant than alanine, free Glu is rela-
tively abundant throughout pregnancy in humans (Table 6.1), whereas in other ani-
mals (i.e., pig), it is the most abundant FAA in both amniotic fluid and maternal
plasma (Wu et al., 1995). From the perspective of early feeding and experiences, the
human fetus actively swallows significant amounts of amniotic fluid, particularly
during the last trimester (Underwood et al., 2005), so the varying amounts of FAAs
and free Glu could result in differential exposures prior to breastfeeding.
6.3.2 Human Milk
For reasons not completely understood, the mammary tissue of many mammals
produces large amounts of nonprotein nitrogenous compounds, including (protein-
unbound) FAAs. Research quantifying the amino acid composition of human milk
dates back to the 1940s and is a growing field of research, perhaps motivated by the
need to improve the composition of infant formula, since human milk is the stan-
dard for human infant nutrition (Ballard & Morrow, 2013).
The total amino acid (both unbound and free) profile and the free Glu content of
human milk at the three stages of lactation, colostrum (0–5 days), transitional milk
(6–20 days), and mature milk (≥21 days), are shown in Table 6.1. Here we highlight
the findings from a systematic review of 22 studies on both preterm and full-term
human milk (Zhang et al., 2013). The inclusion criteria for this meta-analysis were
that the women who donated the human milk samples had to be healthy, have deliv-
ered a term infant of known age, and have to be consuming free-living diets and that
the study provide adequate information for the milk samples, such as stage of lacta-
tion, method of extraction (e.g., type of pump) and storage (e.g., liquid, freeze-dried
form), units of concentration, and geographic location. FAAs had to be quantified
by ion exchange chromatography or an automatic amino acid analyzer and not by
microbiological methods.
The systematic review revealed that total amounts of amino acids (both unbound
and free) decline during the first 4 months of lactation and remain stable thereafter
(Zhang et al., 2013), a pattern that parallels the changing protein needs of growing
infants (Dupont, 2003). From the data in the review (Zhang et al., 2013) and other
publications, we summarized the total FAA content in human milk across lactation
(Table 6.1). The pattern of individual amino acids differs over time. While most
amino acids were significantly lower in mature milk than in colostrum (i.e., leucine,
lysine, phenylalanine, valine, threonine, methionine, isoleucine, taurine, arginine,
asparagine, tyrosine, proline) or stayed the same (i.e., histidine, glycine, serine, cys-
teine, alanine), the FAA glutamine and Glu increased with progressing lactation.
Overall, at each stage of lactation, Glu, glutamine, taurine, and alanine were the
most abundant FAAs, with Glu and glutamine accounting for about half of FAAs at
each stage, a finding consistent with our own research (Baldeon et al., 2014).
To graphically depict the changes in total FAAs and free Glu during lactation, we
focused on our longitudinal study on adolescent and adult mothers that measured
the FAAs including free Glu content in colostrum, transitional milk, and mature
milk over time (Baldeon et al., 2014). Figure 6.1 shows variation in the relative
amount of FAA Glu and all other FAAs in the milk of lactating woman during the
three stages of lactation. What causes this variation and whether variation in the Glu
content of the maternal diet plays a role remain important areas of future research.
Fig. 6.1 Concentration (umol/L) of free amino acid glutamate (FAA Glu; hatched bars) and other
free amino acid concentration (other FAAs; solid bars) in human colostrum (Panel A), transitional
milk (Panel B), and mature milk (Panel C). Each bar represents an individual lactating woman (A,
n = 65; B, n = 47; C, n = 45). The height of each bar represents total FAA concentration. (Data from
Baldeon et al. (2014))
6.3.3 Infant Formulas
For healthy infants fed artificial milks, the four types of infant formula are currently
on the market are cow milk formula (CMF), which is the most commonly consumed
infant formula; soy formula; partially hydrolyzed formula; and extensive protein
hydrolysate formula (EHF), a type of formula typically given to infants who have
cow milk protein allergy or intolerance to intact protein (Rossen et al., 2016).
Although these formulas are isocaloric and can differ in their source of carbohy-
drate (e.g., lactose vs. modified cornstarch), the form of the protein, the degree of
hydrolysis, and in turn the concentration of FAAs are the major distinguishing fac-
tors (Ventura et al., 2012a). For example, the milk proteins (whey and casein) in
hydrolysate formulas are treated with enzymes to break down their protein struc-
ture, generating FAAs, which lessens the burden of digestion and minimizes aller-
genicity for infants (Cook & Sarett, 1982). Thus, while infants fed any of these
formulas are categorized as formula fed, they are not a homogeneous group since
feeding these different formulas will lead to different exposures to proteins and
FAAs, including umami-tasting compounds.
Table 6.1 presents FAA levels for the four types of infant formula. EHF had the
highest levels of FAAs, averaging 120-fold higher than CMF, 39-fold higher than
soy protein formula, and 36-fold higher than partially hydrolyzed formula.
Compared to human milk, the concentrations of FAA and free Glu are lower in
CMF but substantially higher in EHF (Baldeon et al., 2014; Ventura et al., 2012a;
Agostoni et al., 2000a, 2000b). Quality control measures and manufacturing stan-
dards for infant formula minimize variations in FAA content for a given type of
infant formula. For example, infants fed different brands of CMF will have a simi-
larly low exposure to free Glu (Ventura et al., 2012a), compared to those fed breast
milk, and infants fed EHF will ingest substantially higher concentrations of FAAs,
experiencing different tastes (Ventura et al., 2012a). Psychophysical testing among
adult sensory panels has revealed that EHF tastes less sweet, more bitter, and more
savory than CMF, due in part to the differences in FAA content (Mennella &
Beauchamp, 2005; Ventura et al., 2012b).
6.4 Effects of Early Umami Experiences on Taste Acceptance
Before tasting solid foods, infants are exposed to varying amounts of umami tastes
in amniotic fluid, human milk, and the different infant formulas. Mennella,
Beauchamp, and colleagues have used the differential early exposure to free Glu
and umami flavors of infants feed exclusively human milk, CMF, or EHF as an
experimental model system to study early flavor learning, food acceptance, satia-
tion, growth, and the developing microbiome (Ventura & Mennella, 2017; Mennella
& Trabulsi, 2012; Mennella et al., 2009, 2014b, 2022).
Forestell, Mennella, and colleagues (Mennella et al., 2009) used the striking dif-
ferences in the taste and FAA content among human milk and infant formulas to
understand how the earliest feeding experiences modify orofacial reactivity and
intake of umami and other basic tastes both before and after the introduction of solid
foods. At the time mothers decided to add cereal to their diets, 4- to 9-month-old
infants who were exclusively fed human milk, CMF, or EHF were tested on six
occasions, in counterbalanced order, for their acceptance of the basic tastes in a
cereal matrix: sweet (0.56 M D-lactose), salty (0.1 M NaCl), bitter (0.24 M urea),
savory (0.02 M MSG), and sour (0.006 M citric acid).
The type of milk infants were fed with affected their liking and acceptance of the
basic tastes in a food matrix. Breastfed and EHF-fed infants were more likely than
CMF-fed infants to smile (facial relaxation) while eating the Glu-flavored cereal,
which likely reflects their exposure to the high concentrations of free Glu found in
human milk (Baldeon et al., 2014; Agostoni et al., 2000a) and in EHF (Ventura
et al., 2012a). However, breastfed infants do not all have the same flavor experiences
early in life, because human milk has a great degree of individual variation in taste-
reactive chemicals (for review, (Mennella, 2007; Spahn et al., 2019; Mennella et al.,
2017)), including Glu (Baldeon et al., 2014) (see Fig. 6.1), and thus in the taste
experiences of their infants, which presents difficulties in interpreting the results.
After weaning to table foods, as caloric intake from infant formula declined,
infants’ preferences for the basic tastes in cereal reflected the types of foods they
had been fed: infants who ate pasta and other foods that contained cheese or toma-
toes, which have naturally occurring free Glu, showed greater acceptance of the
MSG-flavored cereal (Mennella et al., 2009).These findings are consistent with
results from randomized feeding trials that revealed differences in food preferences
in children fed EHF or CMF years after their last taste of the formula (Sausenthaler
et al., 2010; Mennella & Beauchamp, 2002; Trabulsi & Mennella, 2012): the prefer-
ence for umami flavors depended on the type of milk fed (Mennella & Castor, 2012;
Mennella et al., 2011a). CMF and EHF differ in composition, in the profiles of vola-
tile flavors, and in tastes other than umami, and the evidence suggests that palat-
ability is personal, dependent on experience.
Experimental evidence demonstrates the plasticity in flavor learning during
infancy. By experimentally manipulating the timing and length of exposure to EHF,
Mennella, Beauchamp, and colleagues discovered a sensitive period for flavor pro-
gramming during which feeding EHF renders this formula highly palatable and
accepted throughout infancy (Mennella et al., 2004, 2011a). Infants introduced to
EHF during the first 3.5 months accept its taste, but this acceptance diminishes in
infants first introduced when they are older than 4 months (Mennella et al., 2014b;
Mennella & Beauchamp, 1998). Infants exposed to EHF for at least 1 month during
this sensitive period do not reject its taste when they are older: they feed EHF to
satiation, prefer EHF to CMF, and do not display facial expressions of distaste while
feeding, and mothers perceive their infants enjoy its taste (Ventura et al., 2015;
Mennella et al., 2004, 2011a). Moreover, the flavors experienced during early
breastfeeding and formula feeding “imprint” and remain preferred for a consider-
able time thereafter (Mennella et al., 2009, 2017; Sausenthaler et al., 2010; Schuett
et al., 1980; MacDonald et al., 1994; Owada et al., 2000; Liem & Mennella, 2002;
Hepper et al., 2013). Whether variation in the umami content of the maternal diet
during pregnancy and lactation results in differential exposure by their infants in
amniotic fluid and human milk, respectively, is an important area for future research.
6.5 Satiation, Satiety, and Growth
Relative to intact proteins, hydrolyzed proteins are absorbed and metabolized more
rapidly than intact proteins (San Gabriel et al., 2007). Indeed, in feeding trials,
young infants ingested more CMF to satiation than they did EHF (Ventura et al.,
2012b, 2015; Hyams et al., 1995; Mennella & Beauchamp, 1996; Mennella et al.,
2011b). To our knowledge, only a few experimental studies in infants have investi-
gated the effects of MSG content on satiation (intake within a meal) and satiety (one
meal’s effect on intake at subsequent meal) (Ventura & Mennella, 2017; Ventura
et al., 2012b).
Because Glu is the most abundant FAA in EHF (Ventura et al., 2012a), we con-
ducted a within-subject, two-meal, 3-day study to determine whether the differ-
ences in the satiation effects of CMF and EHF were due to differences in free Glu
content (Ventura & Mennella, 2017; Ventura et al., 2012b). The experimental design
allowed for infants to determine the timing and duration of each meal on each test-
ing day. In randomized order, they were fed one of three isocaloric formulas during
the first meal—CMF, EHF, or CMF—plus added free Glu, in the form of MSG, to
approximate levels in EHF (CMF + Glu). When infants signaled hunger several
hours later, they were fed a second meal of CMF.
As shown in Fig. 6.2, the infants consumed significantly less CMF + Glu and
EHF than CMF during the first formula meal, whereas intake during the second
formula meal did not differ across the three testing days. Thus, adding free Glu to
CMF was sufficient to induce greater satiation compared to CMF alone. That is, the
infants consumed less of the two formulas higher in free glutamate (EHF,
CMF + Glu) (Ventura et al., 2012b) and began displaying satiation behaviors earlier
compared to feeding CMF alone (Ventura et al., 2012b, 2015), and they did not
compensate at the next meal (Ventura et al., 2012b). Thus, levels of free Glu in for-
mula affect intake, suggesting that what infants are fed may be as important as how
they are fed.
As stated earlier, although isocaloric, CMF and EHF differ not only in the con-
tent of FAA Glu but also in macronutrient composition. For example, EHF contains
higher concentrations of FAA and has cornstarch as the carbohydrate source,
whereas CMF contains mainly intact proteins and has lactose as its carbohydrate
source. Nevertheless, long-term feeding trials have consistently revealed more nor-
mative weight gain and decreased risks of diseases during childhood in infants fed
EHF compared to those fed CMF. The growth trajectories of EHF-fed infants were
normative and comparable to infants fed human milk (Mennella et al., 2011b),
whereas infants randomized to CMF experienced accelerated weight gain during
Fig. 6.2 Amount of formula (mL; mean ± SEM) ingested during three separate test sessions that
occurred on three separate days. In counterbalanced order, infants were fed CMF (black bars),
CMF with added Glu (CMF+ Glu; gray bars), or EHF (blue bars) during the first formula meal (A,
Feed 1). Infants were fed CMF alone during the subsequent formula meal (B, Feed 2). Intake dur-
ing the first formula meal, but not the second, differed across the three testing days. Different
superscripts (a, b) indicate significantly different at P < 0.05. (Adapted with permission from
Ventura et al. (2012b))
the first 4 months due to both lower energy loss and greater energy intake (Mennella
et al., 2018). Two of these trials enrolled healthy infants with a family history of
atopy (Roche et al., 1993; Rzehak et al., 2009) for the first 4 months (Rzehak et al.,
2009) or 6 months (Roche et al., 1993), and two other trials both randomized healthy
2-week-old infants with no history of atopy to feed either EHF or CMF for
8.5 months (Mennella et al., 2011b) or 12 months (Mennella et al., 2018). In the
latter trial, while within the range of typically growing infants, body weight-for-
length Z scores between CMF- and EHF-fed infants remained significantly different
during the first year. Because the World Health Organization standards established
the growth of breastfed infants as the norm (WHO Multicentre Growth Reference
Study Group, 2006), that the Z scores of infants fed EHF tracked at zero means their
growth was similar to that of breastfed infants. Three other trials, each of shorter
duration and with fewer subjects, did not report growth differences but did report
greater satiation when infants are fed EHF than when fed CMF (Hyams et al., 1995;
Vandenplas et al., 1993; Borschel et al., 2014).
Taken together, the experimental evidence reviewed herein reveals that infants
respond to differences in the FAA content of their milk, in terms of intake within a
meal in the short term. Whether the higher concentrations of free Glu in human milk
and EHF (see Table 6.1) are the underlying mechanisms for the more normative
weight gain, which is associated with lower risks of obesity in the longer term
(Trabulsi et al., 2020; Zheng et al., 2018), is an important area for future research,
because early rapid weight gain (Mennella et al., 2011b, 2018; Rzehak et al., 2009)
is a consistent and established risk factor for later obesity and other comorbidities
(Zheng et al., 2018; Monteiro & Victora, 2005; Woo Baidal et al., 2016).
6.6 Summary
From an early age, the flavor and metabolic functions of umami taste are evident in
humans. Infants are born with the capacity to detect and prefer umami taste, like
they do with sweet taste, but only when it is presented in the context of a food, not
in plain water, similar to findings in adults. Inherent plasticity is associated with this
taste, which interacts with early experience to vary preference based on what infants
are fed. Beginning with the first foods, the content of free Glu in amniotic fluid var-
ies during pregnancy, as well as during lactation, and varies by type of infant for-
mula. After birth, those fed human milk or protein hydrolysate formulas will have
greater exposure to free Glu than those fed CMF. During early milk feedings, Glu
and perhaps other FAAs can modulate satiation in the short term and growth and
risks of obesity in the long term (Mennella et al., 2014b). We hypothesize that,
because they are unbound, FAAs can be sensed by receptors in both the oral cavity
(Chaudhari et al., 2000b) and intestinal and gastric walls (San Gabriel & Uneyama,
2013), likely conferring beneficial physiologic and metabolic effects (Ventura et al.,
2012b; Mennella et al., 2011b; Burrin et al., 2008).
Experience with umami continues to evolve as children begin to eat the foods of
the table. Like other flavors and tastes, these early feeding experiences “teach” chil-
dren what foods are safe and what foods are eaten and preferred by their caregivers
and family (Mennella et al., 2016b). In general, early experiences change not the
taste quality per se but its palatability—a more labile feature of a food that drives
eating behaviors and food choice (Mennella et al., 2020).
Acknowledgments AS is an employee of the company Ajinomoto Co., Inc. JAM has no conflicts
to disclose. The writing of this manuscript was supported in part by NIH grants R01DC016616,
R03HD94908, and R03HD102303 from the Eunice Kennedy Shriver National Institute of Child
Health and Human Development (JAM). The content is solely the responsibility of the authors and
does not necessarily represent the official views of the National Institutes of Health or Ajinomoto
Co., Inc.
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Zolotarev, V., Khropycheva, R., Uneyama, H., & Torii, K. (2009). Effect of free dietary glutamate
on gastric secretion in dogs. Annals of the New York Academy of Sciences, 1170, 87–90. https://
doi.org/10.1111/j.1749-6632.2009.03900.x
Ana San Gabriel is DVM and MS in Nutrition. She is Science Communicator and Associate
General Manager at the Department of Global Communications in the Ajinomoto Co., Inc. Tokyo,
Japan. Dr. San Gabriel earned her degree in Veterinary Medicine at the Universidad Autonoma of
Barcelona in Spain and MS in Nutrition at the Department of Dairy and Animal Science of the
Pennsylvania State University. She continued her research on lactation regulation at the University
of Tokyo from where she moved to the Research Institute of the current company where she stud-
ied the molecular distribution of umami receptors in tissues.
Julie A. Mennella is a Member at Monell Chemical Senses Center in Philadelphia. She received
her PhD from the University of Chicago in Biopsychology and then conducted her postdoctoral
studies at the Monell Center, where in 1990 she joined its faculty. Over the past three decades, Dr.
Mennella and her colleagues have made significant discoveries on the development and function
of the flavor senses, early nutritional programming, and the effects of alcohol and tobacco use on
women’s health and physiology, publishing more than 200 scientific articles and chapters.
Chapter 7
Umami and Healthy Aging
Minoru Kouzuki and Katsuya Urakami
7.1 Introduction
The World Health Organization has declared 2021–2030 as the Decade of Healthy
Ageing, calling for coordinated actions by governments, civil society, international
agencies, professionals, academia, media, and the private sector over the next
decade to attain sustainable development goals (World Health Organization. UN
Decade of Healthy Ageing, 2020). This includes ensuring that the people, families,
and communities age healthily as the world’s population ages at an accelerating
rate. In Japan, the average life expectancy and healthy life expectancy are among the
highest in the world (World Health Organization. World Health Statistics, 2021),
and this is considered to be due to the high level of medical care and improvement
in the healthcare system relative to the past. However, it has also been suggested that
the Japanese dietary patterns play a role in reducing the mortality risk (Matsuyama
et al., 2021). The typical Japanese diet is a well-balanced diet with a wide variety of
ingredients that leads to the intake of many beneficial nutrients. The Japanese peo-
ple have a great interest in food, and Japan has many of the world’s most advanced
technologies and ideas. Therefore, the contribution of the food industry to health is
likely to be significant. We believe that efforts in various areas are necessary for
healthy longevity, and diet is one of the most important areas.
Humans acquire nutrients predominantly by consuming food, which helps main-
tain homeostasis in the body. In other words, daily diet plays an important role in
maintaining good health throughout one’s life. Especially in older adults, it is
important to maintain good nutritional status by consuming sufficient nutrients
from the diet, not only to prevent diseases but also to delay the functional decline
associated with aging. However, older people are more likely to suffer from
M. Kouzuki (*) · K. Urakami

Tottori University, Tottori, Japan

https://doi.org/10.1007/978-3-031-32692-9_7
148 M. Kouzuki and K. Urakami
malnutrition due to a variety of factors, including decreased food intake, changes in

eating and swallowing functions (oral functions), and changes in taste, which may
occur with aging (Ahmed & Haboubi, 2010; Pilgrim et al., 2015; Landi et al., 2016;
Minakuchi et al., 2018; Iwasaki et al., 2021). They may also suffer from disorders
that result in impaired nutrient absorption and abnormal loss of nutrients from the
body. Malnutrition leads to a decrease in activities of daily living, increased risk of
infectious diseases, prolonged hospitalization, and increased mortality (Lonterman-
Monasch et al., 2013; Marshall et al., 2016; Katona & Katona-Apte, 2008; Hao
et al., 2019; Cederholm et al., 1995). This can have a serious impact not only on the
person but also on the society, because of the possible medical and nursing care costs.
Among the several approaches for improving nutrition in older adults, this chap-
ter focuses on umami taste. L-Glutamic acid and its salt, a type of amino acid, were
discovered in Japan as flavors and flavor enhancers, and its sodium salt, monoso-
dium L-glutamate (MSG), is used as umami seasoning (see also Chap. 1). Umami
taste is accepted as the fifth basic taste, in addition to salty, sweet, sour, and bitter,
and the potential of umami ingredients for enhancing the taste and appetite among
vulnerable individuals has gained attention. Umami for seniors can be discussed
from two points of view: its relationship with appetite, salivation, and taste in older
adults and the effects on nutritional status, quality of life (QOL), and cognitive func-
tion from continuous MSG intake. However, few studies have investigated umami
with older adults as the target group or interventions with MSG in particular.
Therefore, in this chapter, we discuss how umami can be used to improve the health
of older adults by introducing studies that have been conducted not only on older
people but also on the general adult population.
7.2 Umami in Older Adults for Nutritional Health
Undernutrition results from an imbalance between the intake (or absorption) of spe-
cific nutrients and their required amounts, in addition to necessary energy and pro-
tein, and is followed by sequential changes in metabolic function and body
composition. Factors that have been correlated with lower food intake and malnutri-
tion in older adults and are associated with aging are impairment of chewing and
swallowing, compromised digestion and absorption rates, and decline of physical
activity. These factors are also associated with depression, dementia, loneliness, and
isolation (Robinson et al., 2018; Nieuwenhuizen et al., 2010). As a result, health
problems may rise from malnutrition, or malnutrition may cause health problems.
Older people often suffer from some form of disease that may deteriorate their
nutritional status. On the other hand, age-related changes may also lead to problems
with nutrient intake, such as compromised digestion and absorption. This section
considers the following age-related changes: oral function, digestion and absorption
function, dietary intake, and sensory function. Although umami taste alone is not a
“one-size-fits-all” solution for malnutrition, interventions utilizing umami have
reported improved salivary secretion, a factor that causes dysphagia, which affects
7 Umami and Healthy Aging 149
the promotion of digestion, meal enjoyment, and taste sensation, which in turn
relate to appetite and dietary intake. Treating taste disorders may lead to higher
interest in the diet and solve nutritional problems by enabling the sensation of vari-
ous tastes, including umami.
7.2.1 Umami for Food Enjoyment and Appetite Enhancement

Among Older Adults
Improving the enjoyment of eating and appetite is important to prevent malnutrition

and health problems. MSG is a seasoning agent that improves the flavor and palat-
ability of foods. For example, one study investigated whether enhancing flavor
increases appetite (Mathey et al., 2001). One of four flavors, chicken, beef bouillon,
turkey, and lemon butter (fish), was added (1 ± 0.2 g) into the main dish using a
spice shaker to enhance flavor. Each 100 g of added flavor contained about 60 g
sugars/starch and 30 g MSG, as well as protein, fat, salt, and so on. Thus, the added
flavor had a high MSG content. After a 16-week intervention targeting people older
than 65 years living in a nursing home, the body weight of the flavor-enhanced
group increased (mean ± standard deviation: 1.1 ± 1.3 kg) compared with that of the
control group (−0.3 ± 1.6 kg). Daily dietary intake was significantly decreased in
the control group, whereas no significant change was observed in the flavor-
enhanced group, which remained relatively stable. In the flavor-enhanced group, the
consumption of flavor-enhanced cooked meals significantly increased, degree of
daily hunger increased, and the subjective sense of smell improved. Since the sense
of smell plays an important role in meal enjoyment, it is possible that providing the
participants with a meal with a good aroma enhanced their enjoyment of the meal,
which in turn led to increased food intake and weight gain, thereby improving nutri-
tional status. The results suggest a favorable effect of MSG-containing seasonings
on flavor enhancement.
Several studies have examined the effects of adding only MSG. A study that
investigated changes in food palatability, perceived saltiness, and food intake among
young people (18–39 years old) with MSG supplementation in the diet (Bellisle
et al., 1989) found that, depending on the type of meal, the addition of 0.6% MSG
was preferred in spinach mousse, and 0.6% or 1.2% MSG was preferred in beef
jelly. However, as the concentration of MSG increased, the degree of perceived
saltiness also tended to increase, as expected. The addition of 1.2% MSG led to an
increase in dietary intake on the first week of testing, whereas the addition of 0.6%
MSG to the diet increased dietary intake over successive weeks. Thus, high levels
of stimulation may exert rapid effects, but moderate levels may be better for lasting
effects.
Another study, in which 0.6% MSG was added to two lunch menus of older
people to evaluate dietary and nutrient intake in each menu (Bellisle et al., 1991),
found that intake of some but not all enhanced foods increased. This is considered
to result from expecting an appetite-stimulating effect by adding MSG, but there

was no increase in intake of MSG-containing soups with different menus, suggest-
ing MSG may be compatible only with specific foods. Moreover, the influence of
food choices in the diet could also have an effect, as the results were different for
two lunch menus, and intake of calcium and magnesium increased in one of the
menus, and intake of sodium and fats increased in the other. Sodium in MSG is
about one-third that in sodium chloride (NaCl) (Bellisle, 1999), and if the amount
of NaCl is reduced and an appropriate amount of MSG is added, the palatability is
maintained (Morita et al., 2021; Hayabuchi et al., 2020) (see also Chap. 4 Dunteman
and Lee); therefore, the effective use of MSG to reduce salt intake may help prevent
hypertension and even reduce the risk of cardiovascular disease, and from a health
perspective, enhancing flavor and taste with MSG is beneficial. Taken together,
these studies suggest that the use of an appropriate amount of MSG is expected to
increase palatability and appetite, resulting in enhanced enjoyment of eating.
The mechanism of appetite enhancement by MSG may also involve its effect of
promoting the digestion of food. Glutamate is thought to regulate digestive function
not only through receptors in the oral cavity but also through activation of the vagal
afferent fibers from the gastric branch via glutamate receptors in the stomach
(Uneyama et al., 2006; Yamamoto et al., 2009; Toyomasu et al., 2010). A study
conducted in healthy men 27–45 years of age showed that adding 0.5% MSG to
protein-rich liquid meals enhanced gastric emptying compared with the absence of
MSG (Zai et al., 2009), suggesting it is involved in protein digestion. In another
study, targeting healthy individuals 30–50 years of age, MSG or NaCl was added to
lunch and dinner for 7 consecutive days, and then pre- and post-meal assessment
was conducted on day 7. Results showed that MSG supplementation at nutritional
doses elicits elevation of several plasma amino acid concentrations in healthy
humans (Boutry et al., 2011), suggesting that adding MSG may affect uptake of
amino acids in addition to digestion of protein. Adding MSG to the diet may thus
increase dietary intake by promoting digestion and may also improve nutritional
status due to increased nutrient uptake.
7.2.2 Effect of Umami Stimulation on Salivary Secretion

in Older Adults
Dry mouth has been used as a comprehensive term to refer both to xerostomia (the
subjective sensation of dry mouth) and to salivary gland hypofunction (objective
findings of dry mouth), such as hyposalivation or altered salivary components
(Nakagawa, 2016; Thomson, 2015; Han et al., 2015). Decreased salivary secretion
may cause complaints of xerostomia; however, xerostomia may or may not be
accompanied by decreased salivary secretion due to salivary gland hypofunction
(Hopcraft & Tan, 2010; Napeñas et al., 2009). Reduced salivary secretion negatively
affects oral health, and xerostomia affects QOL.
The proportion of patients with xerostomia and salivary gland hypofunction

increases with age, and factors include high prevalence of systemic diseases and
side effects of regular medications (Villa & Abati, 2011; Smidt et al., 2010; Johanson
et al., 2015). Although age-related changes in the structure of salivary gland tissue
have been shown (Moreira et al., 2006), many believe that aging itself does not
affect salivary secretion (Hopcraft & Tan, 2010; Smidt et al., 2010). Dry mouth
causes problems related to food intake, such as inability to chew food thoroughly,
inability to form a bolus, and inability to swallow; therefore, this condition has a
strong relation to nutritional disorders. There is a report that treating xerostomia in
the context of systemic diseases enabled and improved the sensation of umami taste,
improved appetite and body weight, promoted the enjoyment of eating, and
improved health conditions (Satoh-Kuriwada et al., 2012a). Treatment of dry mouth
is crucial considering its role in nutrition in older adults.
Saliva is secreted by the major (parotid, submandibular, and sublingual) and
minor salivary glands. Most saliva secreted into the oral cavity originates from the
major salivary glands. However, in some cases, xerostomia may not be accompa-
nied by a decrease in salivary secretion, suggesting the involvement of a minor sali-
vary gland in xerostomia. Previous studies have suggested an association between
complaints of xerostomia and decreased labial minor salivary gland secretion rate,
even in the presence of normal or reduced salivary output throughout the oral cavity
(Eliasson et al., 2009). It has also been reported that people with xerostomia had a
more remarkable decrease in lower labial minor salivary gland secretion than in
chewing-stimulated whole salivary secretion and that lower labial minor salivary
gland secretion measurement had superior sensitivity, negative predictive value, and
diagnostic accuracy for discriminating xerostomia compared to chewing-stimulated
whole salivary secretion measurement (Satoh-Kuriwada et al., 2012b).
In an attempt to treat dry mouth, a study examined the change in the amount of
salivary secretion of major salivary glands and minor salivary glands by taste stimu-
lation (Hodson & Linden, 2006). Eight healthy subjects 18–55 years of age were
tested to determine whether stimulation with the basic five tastes (sweet, salty, sour,
bitter, umami) increased parotid salivary flow. The relative efficacy for eliciting sali-
vation was sour > umami > salty > sweet ≥ bitter. Another study verified the amount
of stimulation by the basic 5 tastes on minor salivary glands using an iodine-starch
filter paper method, in 11 healthy subjects with an average age of 31 years (Sasano
et al., 2014, 2015). The order from the highest to the lowest amount of salivation by
stimulation was umami > sour > salty = sweet = bitter. The salivary reaction evoked
by umami stimulation lasted longer than that of other stimuli. Regarding sour stimu-
lation, which is commonly associated with salivary secretion, a salivary secretion
equivalent to the umami taste stimulation was induced immediately after the stimu-
lation, but the amount decreased following stimulation; therefore, the effect was not
sustained. Furthermore, the increase in total salivary secretion from the major and
minor salivary glands was transient with sour stimulation but persisted longer with
umami stimulation when 24 healthy volunteers were stimulated with sour or umami
(Sasano et al., 2010).
Another study was conducted with ten older adults with a mean age of 69.5 years
where saliva was collected three times at intervals of 30 min after the ingestion of
food (at 0, 30, and 60 min) (Schiffman & Miletic, 1999). Test foods were chicken
broth, onion soup, corn, and carrots, with and without 2.0%, 1.5%, 3.5%, and 2.0%
MSG, respectively. After 30 and 60 min, the secretion rates of secretory immuno-
globulin A (μg/min) after ingesting food containing MSG were high, due to
increased salivary flow, because no significant differences in absolute concentration
were found. Thus, repeated taste stimulation may affect immune function through
increased salivary flow.
A method using kelp stock containing MSG has also been reported as a treatment
for dry mouth using umami (Satoh-Kuriwada & Sasano, 2015). Twenty women
with an average age of 61.9 years who complained of dry mouth were asked to drink
or to gargle kelp stock five to six times a day when they felt dry mouth. Around 80%
of respondents answered that their dry mouth had improved, and 67% said they felt
the effect after 1 month of use. In addition to improvement of dry mouth, respon-
dents also said that the method “improved roughness in the mouth,” “prevented food
clogging which made it easier to swallow,” and so on. Improvement of various
symptoms related to dry mouth was observed; thus, this may be a practical method
to enhance salivation.
To summarize, based on available data, umami stimulation appears to promote
salivary secretion and improve xerostomia in both the major and minor salivary
glands, suggesting that umami stimulation can be used to improve dry mouth.
However, since there are a relatively few studies in older adults, and many of these
include only small numbers of subjects, further examination is needed.
7.2.3 Perception of Umami Taste in Older Adults
It is generally believed that as we age we begin to prefer stronger flavors (the com-
bination of taste, smell, and irritant properties of foods). Indeed, a study comparing
the strength of flavor and palatability of four food items (bouillon, tomato soup,
chocolate custard, and orange lemonade) found that older adults tended to have a
preference for higher flavor concentrations than did younger individuals (de Graaf
et al., 1996). One of the reasons that older people prefer stronger flavors is a change
in taste function. To date, many studies have been conducted on the relationship
between taste thresholds and aging (Liu et al., 2016; Boesveldt et al., 2011; Mojet
et al., 2001; Yoshinaka et al., 2016; Yamauchi et al., 2002; Methven et al., 2012;
Welge-Lüssen et al., 2011). Most researchers have thus come to the conclusion that
taste function decreases with age; however, which taste sensitivities are reduced dif-
fers across studies. For example, on one study, recognition thresholds for sweet,
salty, sour, and bitter tastes were in the normal range, but recognition thresholds for
umami were elevated (Satoh-Kuriwada et al., 2012a). Thus, age-related deteriora-
tion in taste function can be understood to vary, because sensitivity for all tastes is
not lost; rather, the detection and recognition ability for specific taste qualities may
be impaired.
Low salivary volume, low serum zinc, the effect of comorbidities, and prescribed
medications have been pointed out as underlying factors for decreased taste func-
tion in older adults (Sasano et al., 2014; Ikeda et al., 2008; Kinugasa et al., 2020),
with a variety of factors having secondary effects. It may also result from decreased
signaling mechanisms for taste in the brain. Previous studies have shown that peo-
ple with Alzheimer’s disease dementia (ADD), which is more likely to develop at an
older ages, may exhibit a decline of taste sensitivities (Ogawa et al., 2017; Kouzuki
et al., 2020). The cause of the decline is thought to be due not to impaired transmis-
sion from peripheral receptors but to a decrease in taste-perception cognitive ability
that accompanies brain atrophy and neurodegeneration. In our study (Kouzuki et al.,
2020), many participants, not only those with ADD but also nondementia controls
(NDCs), could not recognize umami. With respect to umami, the cumulative distri-
bution curves for detection and recognition thresholds, for the percentage of correct
answers for each taste solution, differed from those of other taste solutions, espe-
cially with respect to recognition: a concentration higher than that of other taste
solutions was required (Fig. 7.1). However, 21.4% of NDCs were not able to recog-
nize of umami taste even at the highest concentration, and its recognition became
worse with age.
Studies in humans do not necessarily achieve consistent results, due to differ-
ences in the background factors and living environments of the subjects, differences
in the concentrations of taste solutions between studies, and the fact that some
Sweet- DT Salty- DT Umami- DT Sour- DT Bitter- DT

100 100 100 100 100
80 80 80 80 80
%of subject s
%of subject s
%of subject s
%of subject s
%of subject s
60 60 60 60 60
40 40 40 40 40
20 20 20 20 20
0 0 0 0 0
2 4 6 8 10 12 B 2 4 6 8 10 12 B 2 4 6 8 10 12 B 2 4 6 8 10 12 B 2 4 6 8 10 12 B
Concent rat ion st eps Concent rat ion st eps Concent rat ion st eps Concent rat ion st eps Concent rat ion st eps
Sweet- RT Salty- RT Umami- RT Sour- RT Bitter- RT

100 100 100 100 100
80 80 80 80 80
%of subject s
%of subject s
%of subject s
%of subject s
%of subject s
60 60 60 60 60
40 40 40 40 40
20 20 20 20 20
0 0 0 0 0
2 4 6 8 10 12 B 2 4 6 8 10 12 B 2 4 6 8 10 12 B 2 4 6 8 10 12 B 2 4 6 8 10 12 B
Concent rat ion st eps Concent rat ion st eps Concent rat ion st eps Concent rat ion st eps Concent rat ion st eps
NDC ADD
Fig. 7.1 Cumulative curves for detection thresholds (DT, top row) and recognition thresholds (RT,
bottom row) in patients with Alzheimer’s disease dementia (ADD) and in nondementia controls
(NDCs). The taste functions of patients with ADD and of NDCs were evaluated in detail by the
whole-mouth gustatory test using taste solutions for sweet, salty, sour, bitter, and umami, each
diluted to 13 levels. If the participants could not detect or recognize a taste, even at the highest
concentration, those results are indicated as “burst” (B) on the x-axis. (Modified from reference
Kouzuki et al. (2020), licensed under a Creative Commons Attribution 4.0 International License
(https://creativecommons.org/licenses/by/4.0/))
studies evaluate only detection or recognition thresholds. However, these thresholds

do not usually decrease, at least in older adults. Because elevated thresholds imply
the need for more intense taste stimuli for taste detection, it can be understood that
older adults prefer a stronger-tasting diet due to a lower perception of taste.
Reports on improvements to taste function indicate that taking the zinc agent
polaprezinc at 150 mg/day (administered as 75 mg twice daily; 75 mg of polaprez-
inc contains approximately 17 mg of zinc) resulted in improvements in the mean
recognition thresholds for sweet, salty, and sour tastes in 74% of older adults with
taste disorders (Ikeda et al., 2008). Regarding umami, seven cases 62–78 years of
age with reduced umami sensitivity were treated for xerostomia in addition to sys-
temic disease and improved their recognition threshold for umami, as assessed by a
filter paper disk test (Satoh-Kuriwada et al., 2012a). In another study, in 28 patients
45–78 years of age, with complaints of taste impairment, clinical examinations
(blood tests, salivary tests, an oral candida culture test, and oral hygiene tests) and
investigation of systemic diseases and drug prescriptions were carried out, and
appropriate treatment was performed based on these results. After treatment, all
patients showed lower recognition thresholds of umami than before treatment, indi-
cating that loss of umami taste sensitivity can be improved with appropriate treat-
ment (Satoh-Kuriwada et al., 2014). Evaluation of taste function and treatment for
taste disorders are important because decreased interest in meals due to reduced
taste function may reduce appetite and adversely affect nutritional status.
7.3 Clinical Trials for Continued Ingestion of MSG

in Old Age
Here, we introduce the effects of long-term ingestion of MSG in older adults. MSG
transmits gustatory signals to the brain via oral and gastric receptors, affecting
digestive functions by increasing the secretion of saliva (Hodson & Linden, 2006;
Sasano et al., 2014; Sasano et al., 2015) and gastric juices (Zolotarev et al., 2009)
and promoting digestion (Zai et al., 2009; Boutry et al., 2011). There have also been
reports that the neural organization of the primary gustatory cortex receives inputs
from glutamate receptors on the tongue (Schoenfeld et al., 2004) and that umami-
stimulated activation of the primary gustatory cortex (insular and opercular regions)
and orbitofrontal cortex were observed in functional MRI (de Araujo et al., 2003),
suggesting that ingestion of MSG may affect the brain. Although very few interven-
tional studies have involved MSG consumption for a long time by older adults, this
section introduces studies that attempted to improve the nutritional status and QOL
of older adults by MSG intake (Toyama et al., 2008; Tomoe et al., 2009), as well as
those that examined the effects of continuous transduction of taste signals to the
brain by MSG intake on cognitive function (Kouzuki et al., 2019).
7.3.1 Umami and Improved Behaviors and Nutritional Status

in Old Age
Here, we present a study evaluating the improvement of nutritional status and QOL
of older people by long-term consumption of MSG (Toyama et al., 2008; Tomoe
et al., 2009) and provide our opinion on the subject. Older individuals with malnu-
trition are more likely to have reduced QOL, while interventions that improve nutri-
tional status lead to significant improvements in physical and mental aspects of
QOL (Rasheed & Woods, 2013), and nutritional status of older people is a modifi-
able factor associated with QOL.
A study of 11 inpatients (mean age ± standard deviation: 85.8 ± 8.2 years) who
consumed 0.5% (w/w) MSG added to their staple rice gruel three times daily for
2 months reported no change in body weight before and after the intervention and
no change in serum total protein or albumin, an indicator of the nutritional status
(Toyama et al., 2008). However, the number of lymphocytes in blood increased
significantly during the intervention period and then decreased significantly 1 month
after the intervention period. Low lymphocyte count is an indicator of loss of
immune defenses caused by malnutrition (Ignacio de Ulíbarri et al., 2005) and is
affected by increases or decreases in nutritional status. This parameter is perhaps
connected with protecting the body from infection by enhancing immune function,
indicating that glutamate may activate biological defense systems. Moreover, in the
evaluation of daily performance by the nursing staff, “clear speech,” “cheery facial
expression,” and “eye opening” showed more remarkable improvement, which cor-
related with improvements in QOL. In addition, the revised Hasegawa’s Dementia
Scale (HDS-R), a screening test for cognitive function, showed that five patients
improved, three deteriorated, and three showed no change. These results support the
hypothesis regarding positive effects of MSG intake on cognitive function.
These conclusions were subsequently supported using a similar intervention in a
3-month placebo-controlled, double-blind study (Tomoe et al., 2009). In this inves-
tigation, the group that consumed MSG (MSG group) comprised 14 inpatients
(mean age ± standard deviation: 83.0 ± 8.9 years), and the control group comprised
15 inpatients (84.3 ± 9.6 years). Blood tests revealed no increase in albumin, as in
the previously described study (Toyama et al., 2008), but the ratio of reduced-form
albumin to total albumin, considered an indicator of redox status or quality and
quantity of dietary protein ingestion in the body (Kuwahata et al., 2017; Tabata
et al., 2021; Wada et al., 2020), was increased only in the MSG group, suggesting
an improvement in protein nutritional status. Evaluation of daily performance by
nurses without knowledge of the presence or absence of the intervention indicated
improvement in the MSG group after the intervention, with results comparable to
those of the earlier study (Toyama et al., 2008). On the other hand, no significant
changes were observed in HDS-R scores. However, HDS-R as a screening test for
dementia may have inadequate detection power to assess effects of the intervention.
In addition, to evaluate behavior of patients during the actual diet, the researchers
recorded behavior during meals before and at the end of the intervention and had
both videos evaluated by 13 university students. In the MSG group, activity level,
eye opening (e.g., eating awake), swallowing (e.g., timing of swallowing), cheery
expression, motion of arms and hands (e.g., handling of cutleries), and position
holding showed improvement; overall dietary behavior was improved, and there
was a tendency to try to consume independently.
In summary, these results suggest that continuous intake of MSG in the older
adults may improve the immune system and nutritional status, as evidenced by
improvement in some biochemical markers, and may also contribute to improve-
ment of QOL based on behavioral changes.
7.3.2 Umami and Slower Cognitive Decline in Old Age

with Dementia
Studies to verify brain activation by MSG stimulation have been reported

(Schoenfeld et al., 2004; de Araujo et al., 2003). In addition, as mentioned above,
HDS-R scores improved in 45.5% of patients when MSG was added at 0.5% (w/w)
to rice gruel in each meal given three times a day for 2 months (Toyama et al.,
2008), suggesting MSG may have a beneficial effect on the brain. Therefore, we
investigated the impact of continued MSG consumption on cognitive function and
interest in food in older people with dementia (Kouzuki et al., 2019). The subjects
of this study were 159 dementia older persons living in hospitals or nursing homes
(e.g., geriatric health service facilities, special nursing homes for the aged, and
group homes). The subjects were divided into two groups: one with MSG added to
their daily diet (MSG group) and the other with NaCl added as a placebo (control
group); the dietary intervention was performed for 12 weeks. MSG (0.9 g/dose) or
NaCl (0.26 g/dose, equivalent to the amount of sodium contained in the molecular
content of MSG) was added to three meals daily: breakfast, lunch, and dinner. When
applicable, MSG or NaCl was added to rice porridge, miso soup, or other soup;
otherwise, these additives were mixed in the main dish. After completion of the
intervention, a follow-up period without dietary intervention was provided for an
additional 4 weeks, and examinations of dementia symptomatology, blood tests,
daily performance, and preference for diet were conducted pre- and post-intervention
and post-follow-up.
Cognitive function was tested using the Touch Panel-type Dementia Assessment
Scale (TDAS) (Nihon Kohden Corporation, Tokyo, Japan) (Inoue et al., 2011) as a
subjective method to assess subjects’ cognitive functions. TDAS is a cognitive func-
tion test introduced to touch-panel computers by partially modifying the Alzheimer’s
Disease Assessment Scale, which is considered the most reliable decision-making
method for progress of ADD and treatment effectiveness. TDAS scores did not dif-
fer significantly between the baseline, post-intervention, and post-follow-up groups
Fig. 7.2 The Touch Panel-type Dementia Assessment Scale (TDAS) score: overall (a) and mean
change from the baseline (b). BSL = baseline, INT = intervention, FU = follow-up, Cont = control,
MSG = monosodium L-glutamate. All data represent mean ± standard error; the numbers above the
error bars are the mean values. **p < 0.01. (Modified from reference Kouzuki et al. (2019),
licensed under a Creative Commons Attribution 4.0 International License (https://creativecom-
mons.org/licenses/by/4.0/))
in either the MSG or the control group (Fig. 7.2a); however, comparisons of scores
between the two groups from baseline showed significant improvement in the MSG
group after follow-up (Fig. 7.2b). Thus, it is possible that improved cognitive func-
tion was observed in a test performed at 4 weeks after the intake was discontinued,
thereby indicating the necessity of further investigation. In addition, when examin-
ing the correlation between changes in baseline and post-intervention TDAS scores
and a food palatability survey, we found a significant association between the total
TDAS score and the enjoyment of the meal in the MSG group and a trend toward a
correlation between the total TDAS score and the deliciousness of the meal. In other
words, the greater the improvement in food quality, the greater the improvement in
cognitive function.
Although MSG enhances umami taste, we considered the effect of MSG inges-
tion on taste function. It has previously been reported that the percentage of people
with low serum zinc levels rises with age (Ikeda et al., 2008; Kogirima et al., 2007);
this tendency was also observed in subjects in this study (Kouzuki et al., 2019), with
baseline mean serum zinc levels as low as 61.8 μg/dL in the MSG group and 63.5 μg/
dL in the control group (reference value in the study, 64–111 μg/dL). Zinc plays an
important role in taste bud homoeostasis, and patients with taste disorders have
exhibited significant improvements in taste sensitivity after treatment with a zinc-
containing compound (Ikeda et al., 2008; Sakagami et al., 2009). The variation in
serum zinc levels before the intervention did not significantly differ between the two
groups compared to variation after the intervention and at follow-up, but the MSG
group showed increased serum zinc levels in the post-intervention test. Zinc is
absorbed from the intestine, and the ingestion of MSG increased the secretion of
gastric juice and upper gut motility via vagus nerve stimulation (Toyomasu et al.,
2010; Zai et al., 2009; Boutry et al., 2011; Zolotarev et al., 2009) and enhanced
digestive absorption, which may have led to a better absorption of zinc. In rats, the
average life span of a taste bud cell is about 250 ± 50 h (Beidler & Smallman, 1965).
It was suggested that zinc deficiency induces delayed proliferation of taste bud cells
(Hamano et al., 2006). Although we cannot make a clear conclusion, taste bud
regeneration in response to increased zinc absorption during the MSG intake period
might have appeared as a sustained effect even after MSG discontinuation. These
results suggest that elevation of serum zinc caused the regeneration of taste buds,
and the effect of MSG on the umami receptors T1R1 + T1R3, mGluR1, and
mGluR4 in taste cells (Yasumatsu et al., 2015) led to a greater perception of the
taste of cooked meals and affected cognitive function by enhancing the taste signal-
ing to the brain via glutamate receptors in the oral cavity, as well as via the vagus
nerve from the stomach (Tsurugizawa et al., 2008; Tsurugizawa et al., 2009).
7.4 Conclusions
Life expectancy is increasing worldwide; however, it is important for healthy life

expectancy to increase concomitantly. Preventive measures to avoid becoming ill
are important to maintain health, and this chapter focuses on nutrition as one of the
factors associated with disease. Although age-related changes may lead to a decrease
in nutritional status, a variety of methods have been shown to potentially prevent or
improve malnutrition. Adding substances that elicit umami taste, such as MSG, has
been proposed as a way to address malnutrition. It may also be possible to promote
nutrient intake from meals by increasing interest in meals by enhancing appetite and
by promoting digestion via the addition of MSG, thereby increasing the amount of
salivation by the stimulatory action of MSG and improving the ability to recognize
umami through appropriate treatment targeting decreased taste function. Long-term
consumption of MSG by older adults is also expected to improve nutritional status,
QOL, and cognitive function. One’s daily diet is important for living healthily even
in old age, and we believe that the approach using umami taste has many possibili-
ties for preventing or improving various health disorders by enhancing the palat-
ability of the diet.
Acknowledgments We thank Dr. Hisayuki Uneyama (Ajinomoto Co., Inc.) for his critical review
of the manuscript. In addition, we thank the editors (Dr. Ana San Gabriel, Dr. Gary Beauchamp,
and Tia Rains) for their assistance.
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Minoru Kouzuki is a junior associate professor in the Department of Biological Regulation,

School of Health Science, Faculty of Medicine, Tottori University, Tottori, Japan. He received a
Ph.D. degree in Health Science from Tottori University in 2018. His research interests include
early detection of dementia by clinical examinations, prevention of cognitive decline, and health
promotion for older adults.
Katsuya Urakami is Professor at Tottori University School of Medicine. He received his

Ph.D. from Tottori University School of Medicine after what he became Assistant and consecu-
tively Associate Professor of the Department of Neurology, Tottori University School of Medicine.
In April 2001, Urakami became Professor at the Department of Biological Regulation of Tottori
University School of Medicine. He was later Specially Appointed Professor, Department of
Dementia Prevention, Tottori University School of Medicine. He is currently Director of the
Japanese Psychogeriatric Society and the Japan Gerontological Society and Chairman of the
Japanese Society of Dementia Prevention.
Chapter 8
Umami Taste as a Component of Healthy
Diets
Ana San Gabriel and Tia M. Rains
Suboptimal diets on top of social and environmental challenges have resulted in a

global nutritional crisis (Independent Expert Group, 2021). Current trends of
unhealthy and unsustainable food production, together with an estimated growth in
population to about 10 billion by 2050, represent a threat to food security, the health
of people, and the health of the planet (Pörtner et al., 2022; Willett et al., 2019). For
the shift to sustainable food systems that provide a healthy, nutritious, and sustain-
able diet, experts have evaluated the best available evidence (Afshin et al., 2019)
and identified a need to increase ingestion of plant-based foods and to reduce con-
sumption of animal-based foods (Willett et al., 2019). Therefore, the amount and
kinds of foods we choose to eat have an impact not only on our health but also on
the environment (Springmann et al., 2016).
The challenge is to make consumption of plant-based foods more appealing. The
sense of taste has great relevance to nutrition: it helps identify and support ingestion
of nutrients, improving the probability of survival (Mattes, 2021). In this context,
the pleasant taste conferred by umami compounds could play a key role in increas-
ing the palatability of plant-based foods and, more specifically, local plant-based
crops that are part of the traditional cuisine in each region.
A. S. Gabriel (*)
Global Communications, Ajinomoto Co., Inc, Tokyo, Japan
T. M. Rains
Research and Development, Ajinomoto Health & Nutrition North America, Itasca, IL, USA

https://doi.org/10.1007/978-3-031-32692-9_8
166 A. S. Gabriel and T. M. Rains
8.1 Diet, Health, and the Environment
The Global Burden of Disease Study 2017 showed an association between the quan-
tity and type of foods we eat and our health (Afshin et al., 2019). The relevance of
nutrition for our health has been strongly reinforced in the last 2 years with the
COVID-19 pandemic (Pörtner et al., 2022; Willett et al., 2019): individuals infected
with SARS-CoV-2 that presented age-associated preexisting comorbidities, such as
noncommunicable diseases (NCDs) like diabetes and obesity (Afshin et al., 2019;
Michael A Clark, 2019), had a higher risk of mortality (Antos, 2021; Ssentongo,
2020; Centers for Disease Control and Prevention, 2022). In fact, the pre-COVID-19
prevalence and incidence of these chronic diseases, which are indicators of poor
metabolic health (but not diseases exclusive to modern humans), have increased and
continue to be the leading cause of total lives lost (Thompson, 2013; WHO, 2020a).
In 2019, before the COVID-19 pandemic, seven out of ten deaths were the result of
NCDs, with cardiovascular diseases at the top of the list—the most frequent chronic
illnesses in an aging population (Tubiello et al., 2021).
Suboptimal diets were estimated to pose a higher risk of mortality from NCDs.
Inadequate consumption of fruits, vegetables, nuts, seeds, and whole grains and
excess consumption of sodium have caused 11 million deaths and 255 million
disability-adjusted life-years (DALYs), a combination of years of life lost due to
premature mortality (YLLs) and years of healthy life lost due to living with disabil-
ity (YLDs) (WHO, 2020b). Eating at least 400 grams of fruits and vegetables
(including legumes, nuts, and whole grains), more unsaturated fats, and less than 5
grams of salt per day throughout the life course was found to prevent all forms of
malnutrition and lower the risk of NCDs (WHO, 2020c).
And, it so happens that these foods not only are more beneficial for our health but
also have a lower impact on the environment (Chai, 2019; Clark, 2019; Springmann,
2016; Tilman & Clark, 2014): the composition of the diet strongly affects the emis-
sion of greenhouse gases. For instance, the production of plant-based foods releases
a lower volume of greenhouse gases than does producing animal-based foods.
However, selecting foods solely on their environmental impact may not automati-
cally maximize human health (Tilman & Clark, 2014).
Thus, we have a diet-health-environment dilemma, in which no region of the
world meets the recommended consumption of health-promoting foods, unhealthy
foods are overconsumed, and unsustainable food systems are causing anthropo-
genic changes to Earth’s climate.
8.1.1 Shift of Global Traditional Diets
As nations become more affluent, especially low- and middle-income countries,

traditional diets based on coarse starchy crops with complex carbohydrates (e.g.,
cereals, roots, and tubers) are replaced by refined starches, added sugars, vegetable
8 Umami Taste as a Component of Healthy Diets 167
oils, and animal fats, and plant proteins are replaced by such animal products as
eggs, meats, and dairy (Drewnowski & Poulain, 2018). These changes in dietary
patterns, supported by greater wealth and higher food affordability, are known as
nutrition transition (Drewnowski & Popkin, 1997). However, the health and envi-
ronmental costs seem to be hidden in inexpensive global staple grain and sugar
crops, as shown by the increase in the burden of malnutrition (Fanzo, 2018; Tilman
& Clark, 2014). A “Western/unhealthy” pattern has been described as having a high
intake of refined grains and sweets, meat, and soft drinks (Murakami et al., 2018).
8.1.2 Umami and the Japanese Dietary Pattern
According to the WHO (2019), Japan has the longest average life expectancy in the
world, which has been partially attributed to the Japanese dietary pattern. Assessed
as the Japanese Dietary Index, the Japanese dietary pattern has been associated with
a lower risk of all causes of death and of cardiovascular and heart disease mortality
(Abe et al., 2020; Matsuyama et al., 2021). A higher adherence to this diet was
associated with a longer life and longer disability-free survival (Abe et al., 2020;
Zhang et al., 2019). Experts have known for some time that evaluating dietary pat-
terns, in which foods are eaten in combination, rather than listing foods in isolation,
gives better guidance for diet quality (Reedy et al., 2014). Common foods in the
Japanese dietary pattern, such as seaweeds, fish, green and yellow vegetables, and
green tea, contain myriad beneficial nutrients and phytochemicals, which are sus-
pected to have a cumulative effect.
The Japanese dietary pattern traditionally known as washoku (a traditional
dietary culture of Japan) is not the only one that lowers the risk of all causes of
morbidity and mortality, but it is the only one that includes a specific taste as part of
its traditional heritage: the umami taste. The guiding principles of washoku, desig-
nated a UNESCO intangible cultural heritage in 2013, explain that the basic struc-
ture of a customary Japanese meal includes a distinctive flavor that results from the
combination in the mouth of the taste, smell, and the tactile sensation of each ingre-
dient (Ninomiya, 2016). The core flavor of many Japanese recipes is the umami
taste from dashi soup stock. The extraction of umami substances when preparing
the stock from traditional ingredients—dried kelp, dried bonito, or dried shiitake
mushrooms—in combination with such products as vinegar, miso, or soy sauce
intensifies the flavor of seasonal and fresh local ingredients (Kumakura, 2015). The
style of eating small portions of a large variety of seasonal foods, including fish and
abundant vegetables, and the effective use of umami taste seem to be the basic ele-
ments that promote positive health outcomes from washoku (San Gabriel et al., 2018).
The modern Japanese diet has transitioned to include high intakes of refined
grains (white rice and white flour), as well as vegetables, seaweeds, soybean prod-
ucts, fish, and green tea, together with low consumption of whole grains, nuts, pro-
cessed meats, and soft drinks (Micha et al., 2015; Murakami et al., 2018). Although
there are still lower rates of morbidity and mortality from coronary artery disease in
Japan than in other regions, this seems to be supported by higher consumption of

plant and marine food and decreased intake of refined carbohydrates and animal fat.
In fact, the Japanese dietary pattern has been changing since the end of World
War II, becoming more diversified and Westernized. The consumption of total fats
and oils has increased threefold from 1960 to 2005, and the consumption of animal
products (meat, poultry, milk, and dairy) increased fourfold, while the intake of
white rice was reduced by half in this period. During these 45 years, the mean intake
of fish and beans augmented slightly, and vegetable consumption remained constant
(Tada et al., 2011). The estimated percentage of energy derived from fat in the
Japanese diet has transitioned from 7.0% in 1946 to 26.6% in 2000, together with a
gradual decrease on salt intake from 13.7 g/day in 1976 to 10.6 g/day in 2006.
Overall, the Westernization of the Japanese dietary style has increased the number
of people with higher atherosclerotic risk by augmenting the rates of obesity, dys-
lipidemia, and hyperinsulinemia with impaired glucose intolerance, especially
among the younger population in Japan (Tada et al., 2011).
Murakami et al. (2018) used principal component analysis to identify three major
dietary patterns through the Japanese National Health and Nutrition Survey from
2003 to 2015: a “plant food and fish” pattern, a “bread and dairy” pattern, and an
“animal food and oil” pattern. They found an apparent continuation of the
Westernization of the Japanese diet: a gradual decrease in consumption of foods in
the plant food and fish pattern and a significant increase in consumption of foods in
the bread and dairy and the animal food and oil patterns (summarized in Table 8.1).
Interestingly, the plant food and fish pattern, which seems to be closer to the recom-
mended “healthy/prudent” pattern, also showed a higher use of salt-based season-
ings (Murakami et al., 2018). Without forgetting the need to decrease the intake of
salt at the population level, experts still express the need to promote a high con-
sumption of plant and fish foods while discouraging the consumption of refined
carbohydrate and animal fat for better health outcomes (Tada et al., 2011). In this
Table 8.1 Dietary patterns in the Japanese population

Dietary Level of
pattern Most consumed foods adherence
Plant food Rice, potatoes, sugar, pulses, green and yellow vegetables, other Low
and fish vegetables, pickled vegetables, fruit, mushrooms, seaweed, fish,
tea, salt-based seasonings
Bread and Bread, sugar, fruit, dairy products Moderate
diary
Animal Other vegetables, red meat, processed meat, eggs, vegetable oil High
food and oil
Adapted from Murakami et al. (2018) based on the Japanese National Health and Nutrition Survey
(2003–2015)
The plant food and fish pattern integrates salt-based seasonings as a food group. Murakami and
colleagues calculated the association of most consumed foods with each dietary pattern by princi-
pal component analysis, using intake of 31 food groups, and scored the level of adherence. We
have simplified this as low, middle, and high adherence to each dietary pattern, with high adher-
ence meaning tested individuals consumed more foods in that category
context, returning to the core flavors of washoku based on the umami taste of tradi-
tional ingredients could help halt the speed of Westernization of the Japanese diet
and move it more toward the plant food and fish pattern while helping reduce excess
salt intake with the use of the umami seasoning (see Chap. 4) (San Gabriel
et al., 2018).
8.2 The Significance of Umami Taste in Food Choice
The sensory cues of foods before, during, and after eating direct our selection of
foods and are at the center of palatability. Smell and taste drive palatability, depend-
ing on our hedonic evaluation of food (Drewnowski, 1997; Gervis et al., 2022;
McCrickerd and and Forde, 2016; Yeomans, 1998). But while orthonasal exposure
to odors through the nose combined with retronasal odors arising from the mouth
provides cues about the food itself in anticipation of eating, taste seems to play a
clearer role in sensing nutrients during and after ingestion of foods (Boesveldt & de
Graaf, 2017). Often cited as the “nutritional gatekeeper” of the body, the sense of
taste has a prominent role in voluntary food ingestion because it helps us choose
what to consume and how efficiently these foods will be digested and metabolized
(Breslin, 2013). In combination with smell and the tactile sensation from the texture
of foods, taste produces flavors and drives a primal response of “acceptable” or
pleasantness, when we detect nutrients that we consider safe to ingest, or “unaccept-
able” or unpleasant, when we estimate that something could be toxic. And this dis-
tinction between toxic and nutritious appears to be qualitatively defined by taste, the
oral perception of chemicals in food (Breslin, 2013). Unique types of taste cells
express specific receptors to detect only one of the five basic tastes: sweet, sour, bit-
ter, salty, or umami (Chandrashekar et al., 2006). The perception of these taste qual-
ities results from the presence in foods of hydrophilic sugars, acids, alkaloids, salts,
and amino acids that dissolve in saliva and activate distinctive receptors in taste
cells, the same receptors that are used to sense molecular changes in our internal
milieu (Bachmanov et al., 2014; Vincis & Fontanini, 2019). The gastrointestinal
tract continues detecting the presence of nutrients and harmful compounds via the
same taste receptors, but in this case food substances evoke not conscious taste sen-
sations but metabolic responses (Breslin, 2013; San Gabriel & Uneyama, 2013;
Steinert & Beglinger, 2011). The brain integrates taste signals from the gustatory
system with other nontaste modalities such as texture, temperature, odor, and even
visceral and homeostatic signals that help us contextualize food experiences (Vincis
& Fontanini, 2019). This is why tastes and flavors that are connected to nutrients
and calories will become more pleasurable over time than those flavors that we
associate with feeling ill from previous experience (Breslin, 2013). The integration
of food signals drives our interpretation of whether a meal is pleasant and ultimately
influences our eating behavior (Khan et al., 2021; Rolls, 2009; Small, 2012).
In the case of umami, the taste of monosodium glutamate (MSG) when presented
alone is not pleasant (Beauchamp, 2009; Okiyama & Beauchamp, 1998), but
functional brain imaging shows that the combination of MSG with a consonant
savory smell such as a vegetable odor induces higher signals of pleasantness in
brain cortical regions where taste and olfactory signals converge. Thus, some
describe umami as a “rich and delicious flavor” (McCabe & Rolls, 2007). In fact,
among the five basic tastes, the savory umami taste of the amino acids glutamate
and aspartate in combination with 5′-ribonucleotides is one of the tastes we perceive
as pleasant in a food context, and these umami compounds are widespread among
many of the foods we eat daily (Breslin, 2013; Ninomiya, 1998).
8.3 The Especially Human Taste of Umami
Umami receptors (described in Chaps. 1 and 2) are the means to sense the proteo-
genic amino acids. However, the high sensitivity of the umami receptor T1R1 + T1R3
for glutamate is very specific to humans. It is hypothesized that in early evolution,
the sense of taste may have helped our ancestor hominids identify nutritious foods
(Breslin, 2013; Toda et al., 2021). Modern humans lean toward the consumption of
more digestible animal proteins instead of plant proteins, but for meats to acquire
strong umami flavors, it is necessary to prepare or ferment them (Mouritsen &
Styrbeak, 2020). Thus, in periods of low food availability, our ancestors may have
had to depend on less digestible and less palatable wild plants (Milton, 2000).
A recent study that combined functional, behavioral, phylogenetic, and ecologi-
cal methods has found that the umami receptor in primates has evolved at least three
times in parallel with the dietary transition from insects to leaves as typical sources
of protein (Toda et al., 2021). The T1R1 + T1R3 receptor in primates whose diet
depends mostly on insects is most potently activated by 5′-ribonucleotides, whereas
in primates whose diet mostly depends on leaves, glutamate evokes the more sig-
nificant response. According to the analysis of Toda et al. (2021), glutamate is one
of the most abundant free amino acids in insects and plants, but plants have lower
5′-ribonucleotide content than do insects. This is because plants do not have muscle
tissue, which is a source of ATP, the precursor for free 5′-ribonucleotides (Mouritsen
& Styrbeak, 2020). Together with evidence from other studies, it seems that the
modern human T1R1 + T1R3 receptor evolved from early mammals to perceive
5′-ribonucleotides present in insect-based diets but later evolved higher sensitivity
to glutamate to facilitate the consumption of leafy plants.
However, plants also contain many bitter metabolites that bind to a family of
mainly broadly tuned bitter taste receptors, the T2R receptors (Adler et al., 2000;
Behrens et al., 2007; Chandrashekar et al., 2000). Because plants contain many bit-
ter metabolites, by considering how bitter taste functions, it is easier to understand
the role of umami taste in modulating the noxious bitterness of leafy greens that are
so important to reduce the risk of diet-related chronic diseases (Afshin et al., 2019;
Drewnowski & Gomez-Carneros, 2000).
8.3.1 Peculiarities of Bitter Taste
Based on their agonist spectra, human T2Rs can be broadly, intermediately, or nar-
rowly tuned (Behrens & Meyerhof, 2013). Bitter taste receptors are thought to func-
tion as warning sensors that prevent humans from ingesting noxious food molecules,
responding to a vast range of compounds known to be bitter to humans. Twenty-five
bitter taste receptor genes in humans (hTAS2Rs) have been identified that encode
for the G-protein-coupled receptor family of type 2 taste receptors (T2Rs) (Behrens
et al., 2007). They have been described as “chemosensory sentinels” because they
alert us to potential threats and trigger defensive responses in the oral cavity and
beyond (Harmon et al., 2021). Bitter taste receptors are also found in chemosensory
cells of extraoral tissues such as the gut and airways that mediate responses from
ingested or inhaled substances (Deshpande et al., 2010; Wooding & Ramirez, 2022).
T2Rs are preeminent in the rejection of potentially harmful compounds that at high
levels can be harmful and even fatal, making us avoid intensely bitter-tasting toxins.
This is why humans normally react by spitting, evading, or vomiting strong bitter
substances (Breslin, 2013). However, many bitter compounds at low levels have
medicinal properties (Bayer et al., 2021). Therefore, compounds that activate T2Rs
are structurally diverse and include different drugs that can be potentially toxic
(Mennella et al., 2013).
The work of Meyerhof et al. (2010) indicates that the perception of most bitter
compounds is not a simple association between an agonist and a specific receptor
but, rather, a complex interaction with a wide set of TAS2Rs (Wooding et al., 2021).
In addition, there are large individual differences in the sensitivity for bitter taste
compounds. The oldest known example refers to the genetic ability to taste two bit-
ter compounds, phenylthiocarbamide (PTC) and 6-n-propylthiouracil (PROP)
(Tepper et al., 2009). Around 70–75% of individuals worldwide taste PTC and
PROP as moderately to intensely bitter—they are considered “tasters”—whereas
the rest (around 28%) are “nontasters”: they are taste-blind to both compounds
(Kim & Drayna, 2004). Psychophysical studies have shown that the population can
be divided into three distinctive groups: nontasters (30%), medium tasters (50%),
and supertasters (20%) (Bartoshuk et al., 1994). This is explained by three single-
nucleotide polymorphisms of the human gene TAS2R38 that cause substitution of
three amino acids at positions P49A, A262V, and V296I. These substitutions pro-
duce two common haplotypes, the taster variant (PAV) and the nontaster variant
(AVI) (Tepper et al., 2009). PROP-sensitive individuals are carriers of the dominant
allele PAV/PAV (the supertasters) or PAV/AVI (the medium tasters), whereas PROP-
insensitive individuals are carriers of the recessive allele AVI/AVI. Others have
shown that the relative expression level for PAV TASR38 mRNA among heterozy-
gous individuals differs widely and correlates with how they rate the bitterness
intensity of both PROP and broccoli juice, which contains PTC-like glucosinolates
(Lipchock et al., 2013).
These differences in the hTAS2R38 gene affect nutritional and health outcomes.
The hTAS2R38 protein is considered the antithyroid-toxin receptor—it detects
thiourea moieties such as PTC-like glucosinolates, which are dietary goitrogens

present in many vegetables, including broccoli. One theory is that sensitivity to the
bitterness of PROP has been conserved as a mechanism to protect humans against
the excessive consumption of dietary goitrogens, especially in environments with
low-iodine soils, where ingestion of plant glucosinolates may aggravate endemic
goiter (Sandell & Breslin, 2006). Infants and children are thought to be more sensi-
tive to the effect of glucosinolates, which may explain their innate aversion to bit-
terness, to prevent the risk of consuming toxic compounds (Breslin, 2013; Mennella
& Bobowski, 2015).
Many cruciferous plants of the family Brassicaceae, including such species as
broccoli, Brussels sprouts, cabbage, cauliflower, watercress, radish, kale, and mus-
tard, contain PTC-like glucosinolates (Sandell & Breslin, 2006). The amount of
these bioactive compounds present in these vegetables has been shown to be a major
barrier for acceptance and consumption of cruciferous vegetables (Drewnowski &
Gomez-Carneros, 2000). However, PROP tasters rated as unpleasant not only the
taste of cruciferous cultivated vegetables but also tastes of other foods like coffee
and grapefruit. Therefore, bitter taste sensitivity to PROP seems to generalize to
other bitter compounds, which may influence the foods we choose to eat. And as a
result of food preference, variations of TAS2R38 appear to be related to adiposity
and vegetable consumption (Duffy et al., 2010; Tepper et al., 2009).
However, not all substances that are toxic taste bitter, nor are all bitter com-
pounds toxic. In fact, associations between bitter taste sensitivity and various health
measures, such as tobacco use, body mass index, glucose homeostasis, and suscep-
tibility to respiratory infections, have also been found (Dotson et al., 2008; Lee
et al., 2012). And it seems that bitter taste acceptance is a behavior we can learn, so
taste education could play an important role in children’s future health (Beckerman
et al., 2017; Mennella et al., 2016).
8.3.2 Interaction of Umami Compounds and Bitter Receptors
In the case of umami taste, human psychophysical studies have shown that umami
substances appear to suppress the bitterness of various compounds (Keast & Breslin,
2003; Keast & Breslin, 2002; Kemp & Beauchamp, 1994; Yamaguchi, 1998). In
vitro assays revealed that MSG, inosine 5′-monophosphate (IMP), L-theanine, and
umami peptides behave as antagonists of various bitter taste receptors, such as the
salicin bitter taste receptor hTAS2R16 and the caffeine receptors hTAS2R43 and
hTAS2R46, also confirmed psychophysically (Kim et al., 2015, 2017). But the
potency to suppress bitterness does not seem to correlate to umami intensity, since
each compound interacts differently with bitter taste receptors (Kim et al., 2015;
Rhyu et al., 2020). Another bitter taste receptor significantly inhibited by umami
compounds is hTAS2R14 (Okuno et al., 2020), the receptor activated by the tea
catechins (−)-epigallocatechin gallate and (−)-epicatechin gallate, which are partly

responsible for the bitterness of tea (Yamazaki et al., 2013), and by several phenolic
compounds from extravirgin olive oil (Cui et al., 2021), whose bitterness indicates
the presence of phenolic compounds.
Altogether, it seems that umami compounds can block the activation of some
bitter taste receptors, but its efficiency is complicated, partially due to the complex
interaction of bitter compounds with TAS2Rs, due to either the broadly tuned (pro-
miscuous) bitter receptors or the intermediately or narrowly tuned (selective) recep-
tors (Bayer et al., 2021; Behrens & Meyerhof, 2013). As shown in Fig. 8.1, the
index of promiscuity (number of substances that activate the receptor divided by the
total compounds of their data set of food TAS2R agonists) is high for TAS2R14,
TAS2R1, TAS2R46, and TAS2R4 and low for TAS2R31, TAS2R20, and TAS2R50.
This means that the ability of umami compounds to block the activation of TASR14
or TAS2R46 may not seem specific, whereas umami compounds are probably more
selective antagonists for that more narrowly tuned receptors TAS2R43 and
TAS2R16. If umami compounds could antagonize the activation of most of the
promiscuous TAS2Rs, we could say that umami inhibits the bitterness of most of
the bitterness of the food components found in plants. However, as yet there are not
enough molecular studies of other broadly tuned TAS2Rs to assume that this is
the case.
Fig. 8.1 Index of bitter taste receptor promiscuity estimated by the number of food agonists to an
individual TAS2R divided by the total number of compounds in the data set. (Modified from Bayer
et al. (2021). Asterisks indicate the bitter receptors for which umami compounds were able to sup-
press the bitterness signal in in vitro assays)
8.4 The Function of Umami for the Consumption of Plants
The importance of consuming vegetables relies on the growing evidence that vege-
tables provide not only nutrients and dietary fiber but also phytochemicals (nonnu-
tritive bitter phenolic compounds) that exert various physiological mechanisms,
have antioxidant activities, and modulate the immune system (Bayram et al., 2018;
Drewnowski & Gomez-Carneros, 2000). These nonnutritive organic constituents,
which are very diverse structurally—including phenolic compounds, carotenoids,
glucosinolates, alkaloids, terpenes, and peptides—are thought to significantly con-
tribute to the health benefit of plant-based diets (Bayer et al., 2021; Cicero et al.,
2017). Yet these are also the molecules that give the astringent and bitter sensory
attributes of vegetables by binding to the human T2R bitter taste receptors (Soares
et al., 2018; Sterneder et al., 2021) and the reason that vegetables elicit complex
perceptual stimuli (Duffy et al., 2010).
8.4.1 Umami, Salt, and Vegetable Consumption
Few sensory studies to date have evaluated the effects of umami on the perception
of bitterness in foods and mixed meals, despite anecdotal evidence suggesting a
benefit. As indicated in Chap. 4, there is a growing understanding of the impact of
umami and specifically MSG on consumer liking and acceptance of reduced-sodium
foods. Studies in that context provide some indication, at least directionally, that
activating the umami receptor via the presence of glutamate attenuates the percep-
tion of bitterness within a mixed meal. For example, Halim et al. (2020) presented
participants with three versions—normal salt, reduced salt, reduced salt with
MSG—of four different mixed meals: roasted vegetables (carrots and eggplant),
quinoa, yogurt-based dip, and pork cauliflower (with onion, peas, and carrots). The
study sought to test the hypothesis that, with addition of MSG, reduced-salt versions
of mixed dishes would be equally as liked as full-salt versions of the same dishes.
In addition to assessing overall liking and liking of appearance, flavor, and
texture/mouthfeel, several sensory characteristics were evaluated by a check-all-
that-apply scale, which included “bitter” in addition to “deliciousness,” “flavorful,”
“balanced,” “bland,” “rancid,” “fresh,” and “savory.” Results showed that, direction-
ally, the attribute of “bitter” was lower in the MSG version of the roasted vegetables,
quinoa, and pork cauliflower dishes. Penalty analysis showed that bitter was a nega-
tive driver of liking overall; therefore, a shift in less perception of bitter may lead to
higher liking.
However, lower bitter with MSG was shown only in the quinoa and the yogurt
dip, where overall liking was higher with the MSG versions than with the standard
full-salt versions. There were many limitations in this study, most notably that the
dishes were not optimized for salt and MSG, and neither dish was considered
Fig. 8.2 A 1952 magazine advertisement for Ac’cent seasoning (monosodium glutamate) from
the United States
particularly bitter. Yet, these results suggest that, in a practical setting, umami may
favorably affect the perception of bitter, which may relate to acceptance of foods.
Additional research, particularly in plant-based foods rich in bitter compounds, is
warranted to better understand the relationship between umami and bitter (Fig. 8.2).
8.4.2 Umami Perception and Dietary Patterns
Various studies have examined the association between umami taste sensitivity and
food preference and/or food consumption, with mixed results (Fluitman et al., 2021;
Gervis et al., 2022; Puputti et al., 2019). Puputti et al. (2019) found that those with
a higher sensitivity to umami consumed more vegetables than those least sensitive
to umami. Fluitman and colleagues found that poor umami taste sensitivity related
to lower adherence to the Mediterranean diet, which is abundant on foods rich in
umami compounds (Fluitman et al., 2021; Ninomiya, 1998). However, in the cohort
study from Gervis et al. (2022), which defined profiles of the perception of all five
basic tastes collectively, older adults with high sensitivity to all tastes but umami
had a higher probability of following a vegetable-rich diet than those with low bitter
or higher umami perception. The difference could be explained by differences in the
cohorts and the type of taste measurements (perception vs. sensitivity). But in the
end, umami is implicated in specific dietary patterns, in addition to individual and
age differences, so it seems important to clarify the association between umami and
overall taste perception and food consumption.
8.5 Conclusion
Taste is often a forgotten element in a sustainable diet, yet it is one of the key drivers
for promoting consumption of a nutritious and healthful dietary pattern (Fig. 8.3).
Of the five basic tastes, umami in particular is often omitted in discussions toward
Fig. 8.3 Steps involved in food preference and food selection, which through the bitter com-
pounds in vegetables ultimately influences cardiovascular health. Umami substances can modulate
various steps by suppressing the signaling of bitter receptors, modifying taste perception, and
increasing food preference, thus helping regulate food choice and vegetable intake. The effective-
ness of umami compounds varies with individual differences and with age: children tend to be
more sensitive to the bitterness of food, and some older adults may lose the ability to taste some
bitter compounds
sustainable food systems that provide healthy, nutritious, and sustainable diets that
better promote health for both people and planet. The global population continues
to grow and age, increasing the prevalence of such noncommunicable diseases
(NCDs) as diabetes and obesity, primary causes of disability and premature deaths
worldwide that result from preventable lifestyle and dietary habits. Psychophysical,
molecular, and nutritional studies continue to show how umami can block the bitter-
ness of many phytochemicals that are cardioprotective and how umami perception
is involved in the preference of healthier dietary patterns. Application of umami
compounds, as has been done for centuries in the traditional Japanese diet, can serve
as a tool to reduce salt and increase the intake of fresh and local plant-based foods.
This aligns with the current trend to increase the ingestion of more sustainable food
choices while reducing consumption of animal-based foods.
Acknowledgments A.S. is an employee of Ajinomoto Co., Inc., and T.M.R. is an employee of

Ajinomoto Health and Nutrition North America, Inc. The writing of this chapter was supported by
the nonprofit International Glutamate Information Service (IGIS). The content does not necessarily
represent the opinions of IGIS or the views of the companies but, rather, is the responsibility of the
authors.
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Ana San Gabriel is Science Communicator and Associate General Manager in the Department
of Global Communications at Ajinomoto Co., Inc., Tokyo, Japan. She earned her DVM at the
Universidad Autónoma de Barcelona in Spain and MS in Nutrition at the Department of Dairy and
Animal Science, Pennsylvania State University. She continued her research on lactation regulation
at the University of Tokyo and then moved to Ajinomoto’s Research Institute, where she studied
molecular distribution of umami receptors in tissues.
Tia M. Rains is a member of Ajinomoto Health and Nutrition North America, Inc., Illinois,
USA. She earned her PhD in Nutritional Sciences from the University of Illinois, Urbana-
Champaign, and her BS in Food and Nutrition from Arizona State University. She has spent over
two decades working in the food and beverage industry in research and development and scientific
communications.
Chapter 9
Practicalities from Culinology®: How
Umami Can Contribute to Culinary Arts
and Sciences
Chris Koetke, Lauren Miller, and Jonathan Deutsch
9.1 What Is Culinology®? What Is Umami?
This chapter differs from the preceding ones by focusing on umami from a practical
gustatory and culinary standpoint, rather than a scientific one. It focuses on the
umami experience and the traditional role that umami plays in foods loved around
the globe. Umami had its place in the culinary world millennia before cooks and
chefs knew what it was scientifically. The day-to-day use of umami in the kitchen
predated and engendered the discoveries of umami as we currently understand it.
Chefs today have the possibility of gaining a sophisticated understanding of umami,
allowing a deeper understanding of the chemical properties of food, from harvest
through eating, and providing tools to make food even more delicious.
This chapter, while not focused on chemistry, includes brief discussions of the
scientific discoveries around umami and the synergistic interaction with nucleotides
(these topics are treated in more detail in Chaps. 2 and 3 of this book). This knowl-
edge reinforces and improves daily sound cooking practices. The intersection of
culinary arts and food science is the locus that gives rise to Culinology®, or culinary
science. The two disciplines inform and rely on each other. This chapter is about the
intentional incorporation of umami to create balanced dishes, whether this comes
from foods intrinsically rich in umami, foods manipulated by further processing to
further develop umami (e.g., fermentation), or the addition of MSG (monosodium
glutamate), the purest form of umami available to everyday cooks. The objective of
this chapter is to apply the science of umami to daily culinary preparation.
C. Koetke (*)
Ajinomoto Health & Nutrition North America, Itasca, IL, USA
L. Miller · J. Deutsch
Drexel University, Philadelphia, PA, USA

https://doi.org/10.1007/978-3-031-32692-9_9
184 C. Koetke et al.
9.2 Umami History Through the Culinary Lens
Jean Anthelme Brillat-Savarin, author of La Physiologie du Goût (Brillat-Savarin,

1826, 1848), written in the early nineteenth century, is most famous for his apho-
rism, “Tell me what you eat, and I shall tell you what you are,” often shortened to
“You are what you eat.” Brillat-Savarin was among the first in Western literature to
document the taste we now call umami, dubbing it osmazome, which Brillat-Savarin
defined as a water-soluble substance in meat containing all its flavors. The term was
used to describe the savory taste indicative of umami. At the time osmazome was
coined, there was no word in the French language to adequately describe the taste
that Brillat-Savarin was experiencing. Not until more than a century later, as the
result of scientific inquiry, did Brillat-Savarin’s taste impressions become under-
stood as umami.
In 1908 Kikunae Ikeda, a professor at the School of Science’s Department of
Chemistry at the Japan’s Imperial University (now called the University of Tokyo),
discovered that MSG was the umami in a broth of kelp seaweed (Lindemann et al.,
2002). Ikeda was the first to identify glutamic acid or glutamate as the source of the
savory taste Brillat-Savarin identified. He named this taste umami and assigned it
the fifth of the five basic tastes. His discovery began in the culinary realm as he
sought to better understand the characteristic tastes in his wife’s dashi, a stock made
from kombu (kelp) and katsuobushi (cooked, smoked, dried, mold-cured, and
shaved bonito) that is foundational to many Japanese dishes. Ikeda used laboratory
facilities at the university to conduct experiments aimed at the extraction of the
umami factor from kelp. Ikeda found that glutamic acid was a central element in the
taste of dashi. Ikeda further recognized that umami was central to many of the foods
he had eaten during a stay in Europe, including tomatoes, asparagus, meat, and
cheese (Yamaguchi & Ninomiya, 2009).
Based on scientific studies on the umami taste receptor on the tongue, published
after 2000, proof of the existence of glutamate receptors has made it widely accepted
that umami is a basic taste (Chaudhari et al., 2000), along with sweet, sour, salty,
and bitter. It is important to note that taste here is limited to the receptor reaction and
different from flavor, which is a broader construct that includes the perception not
only of taste but mainly aroma and also feelings such as heat or cooling. For chefs,
this understanding of glutamate receptors crystalized scientific conclusion under-
scored what they already knew but couldn’t precisely explain. Chefs and cooks
around the world have always cooked using umami-rich sources because they were
simply delicious. Now the science caught up to explain the pleasurable umami taste
from a chemosensory perspective.
One prominent example of umami in ancient cooking is that of the cuisine of the
Roman Empire. Key to much of the culinary fare of ancient Rome was the fer-
mented fish sauces used as essential seasonings throughout the culinary spectrum.
The Romans had four fish sauces: garum, liqumen, allec, and muria. Garum was the
principal sauce produced by the hydrolysis of small fish or fish innards and salt
fermented for several months. Allec was the undissolved fish material remaining
9 Practicalities from Culinology®: How Umami Can Contribute to Culinary Arts… 185
from garum production. Liquamen was very similar to garum in production process
and taste. Muria was the solution from osmosis during the salting of whole, gutted
fish or slices of fish meat. At the time, these umami-rich products were just as ubiq-
uitous and popular as wine and olive oil are today (Curtis, 2009).
The dominant free amino acid in fish sauce is glutamate. While we cannot chem-
ically analyze the fish sauces of ancient Rome, we do know that in the current fer-
mented fish sauces of Southeast Asia, produced through similar salting and
fermenting of fish that resulted in garum, glutamate is present at a concentration of
about 1300 mg/100 mL. Fish sauce also has an inherent nutritional value, providing
other amino acids and numerous micronutrients, such as vitamin B-12 (Nakayama
& Kimura, 1998, Otsuka, 1998, Yoshida, 1998).
9.3 Umami in Foods
Umami, despite its Japanese name, is not a Japanese phenomenon. Rather, it is a

central part of the human taste experience that transcends history or geography.
Cooks and chefs around the world gravitate toward umami-rich foods as a source of
deliciousness (Lioe et al., 2010; Hajeb & Jinap, 2015). Science took thousands of
years to explain what exactly this sensation is that we all crave, and chefs and cooks
know so well. Examples of umami-rich foods in global cuisines include lamb daube
(stew) with anchovy in Mediterranean cooking; bagna cauda (warm dipping sauce
of anchovies, garlic, olive oil, and butter) from the northwest of Italy; koji-fermented
soy pastes from Japan, Korea, and China; soy sauces from numerous Asian cuisines;
cooked tomato and parmesan in Italian cooking; dried meat (machaca) in Mexico;
ketchup on a hamburger; and dried shiitake mushrooms in meat broths in Chinese
cooking.
Science has explained that the umami-rich foods enjoyed around the world are so
impactful because of the presence of free glutamate. This amino acid is what our
body interprets as the pleasurable taste umami. When bound to other amino acids,
as is the case in proteins, glutamate is tasteless. However, certain processes, includ-
ing fermentation, aging, ripening, drying, and low, slow cooking, liberate amino
acids from native proteins, increasing the presence of free glutamate (Wijayasekara
& Wansapala, 2017). Throughout the ages, chefs and cooks have used these tools to
make superlative food. Additionally, some vegetables when ripe are more flavorful
partially due to the increase in glutamate during the ripening process. Tomatoes are
a good example of this (Oruna-Concha et al., 2007; Tommonaro et al., 2021).
Fermentation, responsible for the complex and heady flavors in many foods, also
increases umami. Examples of fermented foods that are rich sources of umami
include natto, Southeast Asian fish sauces, oyster sauce, soy sauce, miso, and kim-
chi fermented with seafood. A number of popular fermented products are based on
either 100% fermented soy or a combination of wheat and soy (Lioe et al., 2010).
Glutamic acid is the predominant amino acid in soybean and wheat proteins (Wang
et al., 2018; Hou et al., 2019). During the fermentation process, a large amount of
glutamate is liberated from these proteins, resulting in a significant increase in

umami. A unique mold koji (Asperigillus oryzae) is central to the production of
many of the products listed above. When fermented with koji, which contains the
enzyme glutaminase, glutamine is converted into glutamic acid. This is the tradi-
tional way of making soy sauce and miso, both of which contain high levels of
umami (Otsuka, 1998; Yoshida, 1998; Lioe et al., 2010; Diez-Simon et al., 2020).
Aging in products such as cheeses, meats, and sausages also results in increased
concentrations of glutamate, contributing to umami taste. For example, during the
ripening of cheese, proteins are broken down into smaller peptides and amino acids,
including not only glutamate but also leucine, valine, lysine, phenylalanine, and
valine. These amino acids and peptides contribute to flavor complexity in cheese
(Umami Information Center, 2016, 2021; Zhao et al., 2016). The increased level of
glutamate (Fig. 9.1) corresponds to the heightened presence of umami. Similarly,
large increases in free amino acids occur during the aging process in cured hams and
other dry-cured meats (Córdoba et al., 1994; Zhang et al., 2019; Heres et al., 2021).
Glutamate is one of the most prominent amino acids in these aged and cured meat
products (Yamaguchi & Ninomiya, 2009).
Drying has the potential to concentrate levels of glutamate, thus making dried
foods more impactful from a gustatory standpoint. An example is dry-cured ham,
which combines increased glutamate from the aging process with further concentra-
tion from the drying process (Zhang et al., 2019). Drying mushrooms results in the
Fig. 9.1 Concentration (mg/100 g) of free glutamate in various cheeses with curation times from
4 months (M) up to 10 years (Y) and in fresh and dried tomato (Fuke & Konosu, 1991). Data were
analyzed at the Japan Food Research Laboratories by the Umami Information Center
5′-nucleotide GMP (guanosine 5′-monophosphate), which has a synergistic effect

on umami perception (Kuninaka, 1960; Yamaguchi et al., 1971). 5′-Nucleotides are
present to varying degrees in different mushrooms. When mushrooms are dried or
frozen, the cell walls are damaged, which converts nucleotides into 5′-nucleotides.
The resulting GMP, combined with the mushroom’s inherent glutamate, with gluta-
mate in other foods, and/or with other free amino acids, gives a pleasurable spike in
umami perception (Wijayasekara & Wansapala, 2017). A good example of this is
dried shiitake mushrooms (Umami Information Center, 2016).
9.4 Concentrated Umami Sources and Products
The previous section explored the history and use of foods inherently rich in gluta-
mate that deliver a strong umami taste. Each of those food ingredients also contrib-
utes other basic tastes, aromas, and textures—flavors that can be advantageous in
various recipes. In contrast, this section looks at highly concentrated forms of
umami that, aside from their strong umami character, contribute little or no addi-
tional flavors and are used in low amounts.
9.4.1 Glutamate
The ionic form of L-glutamic acid, glutamate, is an amino acid that is part of many
proteins. In the production of MSG by fermentation, bacteria convert molasses or
starch hydrolysates into L-glutamic acid, the non-neutralized version of MSG that
tastes sour (because of its acid radical). After neutralization with sodium hydroxide,
the compound adopts the form of sodium L-glutamate, and the umami taste becomes
prevalent (Sano, 2009). This is how MSG is produced in its most inexpensive
form—is readily available, dissolves quickly, and has immediate impact on food
flavor (Lindemann et al., 2002). Before the fermentation method was adopted,
L-glutamic acid was produced from wheat gluten, which contains about 25%
L-glutamic acid by weight (Giacometti, 1979). The pH of most foods is close to
neutral, so glutamate is almost fully found in foods in the form of the tasty sodium
salt rather than the sour L-glutamic acid (Kurihara, 2009).
A good average amount for MSG inclusion is about 0.4% by weight—adding
more does not necessarily render the food inedible, like a larger addition of salt
would (Yamaguchi & Takahashi, 1984), but higher levels of MSG do not necessar-
ily yield noticeably more umami character. On a practical level, MSG can be easily
sprinkled onto a wide variety of savory dishes just as salt would customarily be
used. It does not replace salt but is used in conjunction to provide two different taste
sensations, both of which heighten overall flavor. Thus, MSG can provide an imme-
diate and effortless increase in umami (Beauchamp, 2009).
Yet, in the culinary world, umami is not often considered when creating a dish or
evaluating its overall flavor impact. For instance, chefs are trained to instinctively
consider salt levels in savory dishes but rarely have they been taught to consider
umami levels. When judging culinary competitions or scoring students’ dishes in
culinary schools, chefs often have a rubric for level of seasoning, which refers to
just one taste: saltiness. Yet, considering not just saltiness but the umami axis as
well, along with acid, sweet, and bitter, can provide a much more nuanced assess-
ment of balance in tastes and flavors in a dish than simply “needing salt.”
Keeping a container of MSG alongside salt and pepper is a more complete way
of balancing flavor. For both chefs and cooks, it is important to remember that MSG
is not a salt replacer. Although MSG does contain sodium (as its name, monoso-
dium glutamate, reminds us), in fact MSG contains only 12.3% sodium compared
with table salt’s 39.6% sodium. Using MSG as a 1:1 salt substitution would result
in dishes that are undersalted and therefore out of balance. That said, MSG can be
effective in reducing (but not replacing) salt, as we discuss below (Morita
et al., 2021).
Monopotassium glutamate (MPG), the potassium salt of glutamic acid (just like
MSG is the sodium salt of glutamic acid), is produced in a similar manner.
Fermentation produces glutamic acid, which is neutralized not with sodium hydrox-
ide like MSG but with potassium hydroxide, producing MPG. Both products are
very similar in glutamic acid amount.
MPG delivers concentrated umami with an impact much like MSG’s and simi-
larly dissolves quickly and can be used topically. However, while the umami impact
that MSG delivers is immediate, MPG seems to bring a slightly delayed umami
impact that hits the middle of the palate, leading to a more mouth-filling sensation.
It is also an important ingredient in meat analogs, contributing a “serum”-like flavor
note. It is less readily available than MSG and is more expensive (Kochem &
Breslin, 2017; Morita et al., 2020).
9.4.2 Yeast Extracts
Manufacturing of yeast extract starts with specifically identified yeast organisms

that will produce concentrated sources of umami (Chae & In, 2001). Different yeast
extracts can deliver different levels of glutamate, glutathione, or nucleotides,
depending on the strain. A basic, inexpensive yeast extract has low levels of every-
thing and is used for general background savory notes. Specialty yeast extracts are
more potent and deliver specific tastes and flavors, including umami notes (Jo &
Lee, 2008).
A culture medium is used to grow as much yeast as possible. The yeast is then
washed and goes through a lysis process to break the cell walls, either with enzymes
or through a process called autolysis (Breddam & Beenfeldt, 1991). Autolysis uses
high levels of salt to break the yeast apart. Older-generation yeast extracts contained
high levels of sodium, but newer yeast extracts use more modern techniques that can
reduce the final levels of sodium. The cell wall material is then washed off and the
remainder of the material is filtered and dried (Dimopoulosa et al., 2018). Yeast
extract can also be used in a paste format—a common example of this is Vegemite
spread popular in Australia. From a culinary standpoint, it is essential to know
sodium levels in any yeast extract, as this will impact final taste results.
Utilization of yeast extracts in kitchen applications requires exact measurements,
as they are employed in very small quantities (i.e., slightly below or above 0.15%).
As a result, yeast extracts are more applicable in manufacturing settings than in
daily home or foodservice environments. Using MSG is a much easier way to incor-
porate a concentrated umami source (Okiyama & Beauchamp, 1998). Yeast extracts
are hygroscopic: when exposed to air, they will absorb moisture and clump. Yeast
extracts in combination with nucleotides can have extreme impacts on flavor due to
synergies between glutamates and nucleotides, as we describe below. Yeast extracts
also deliver a consistent umami impact across the palate, but its overuse contributes
an objectionable yeast note (Alim et al., 2019). Overall, yeast extract is less of a
pure form of umami compared to MSG and MPG (J. Formanek, personal commu-
nication, September 9, 2021).
9.5 What Is Not Umami
There is a lot of confusion around what umami is, among home cooks and even
among culinary professionals at high levels, who sometimes confuse the specific
taste of umami with complex flavors like the unctuous sweetness of caramelized
onions, the rich fatness of crispy pork belly, or the deep smokiness of Maillard-
browned barbecued burnt brisket ends (Hofmann, 2005; Finot, 2006). While those
foods may have many wonderful characteristics, including fattiness, unctuousness,
smokiness, brown/roasted flavors, crispness, and so on, unless it was specifically
prepared with umami-rich ingredients, like those described above, it probably had
very low levels of umami taste, as determined both by chemical and by sensory
analyses. Delicious flavor, certainly; umami taste, likely not.
There are a few reasons for the confusion around umami. It is a relative new-
comer to the five basic tastes (Lindemann et al., 2002), and many people were not
trained to recognize it when they were young like they did for the other basic tastes.
Even at the professional level, educators have bemoaned the lack of palate training
in formal culinary education (Deutsch, 2018). Without a firm understanding of
umami, professional chefs do not know how to analyze their foods for optimal
umami. To be umami, there must be a significant amount of glutamate that is not
bound to other amino acids as a taste identified by specific umami receptors on the
tongue (and elsewhere) as indicated in the previous section of Umami History
Through the Culinary Lens. The higher the glutamate concentration in a food, the
more umami will be sensed. In most cases, this sensation cannot be produced from
compounds other than proteins that have glutamate—if no protein is available from
which to extract the glutamate, then there can be no umami sensation on the taste
receptors. If umami receptors are not activated, food may present a delicious flavor
without umami (understanding flavor as the combination of taste with all the other
sensations that influence food perception, such as aroma, texture, juiciness, mouth-
feel, or color) (Grabenhorst et al., 2008). So, there is additional confusion around
ferments that are not protein rich. For example, cabbage is a vegetable low in pro-
tein. Kimchi recipes often add anchovies to cabbage in the fermentation, yielding an
umami-rich product. Without the anchovy, or the production of glutamate by lactic
acid bacteria (Yoon et al., 2021), Kimchi may be salty, sour, and delicious, but it will
not be as rich in umami (Lee et al., 2021).
9.6 Synergies Between Ribonucleotides and Umami
Umami taste can be enhanced through synergistic interactions with the 5′-ribonu-
cleotides GMP and IMP (Yamaguchi, 1998a, b) (see also Chaps. 2 and 3 of this
book). IMP is found in large amounts in meat, poultry, seafood, and dairy, and gua-
nylate is found most significantly in dried mushrooms. Both are also available in
pure commercial/concentrate form (Wang et al., 2020). A small amount of inosinate
and/or guanylate in combination with glutamate creates a strong umami taste (see
Table 9.1).
Chefs have turned to these enhancement combinations (glutamate + inosinate/
guanylate) intuitively throughout history to make food more delicious. Traditional
preparations that employ this synergistic combination include the cheeseburger,
Table 9.1 Umami taste intensity of MSG + IMP mixtures of various percentages
% MSG % IMP Rated taste intensity
100 0 0
95 5 3.5
82 18 6.8
70 30 7.4a
50 50 7.5
30 70 7.3
10 90 5.3
5 95 3.7
2 98 1.8
0 100 0.3b
Data show responses (on a scale of 1 to 10) by trained panelists to the taste of various percent
mixtures of MSG (monosodium glutamate) and IMP (5′-inosine monophosphate), generated by
combining constant concentrations of 0.05 g/dL. Simplified from Yamaguchi (1967)
a
As the percentage of MSG decreases from 100% to 70% and the percentage of IMP correspond-
ingly increases from 0% to 30%, the taste intensity increases. A maximum is reached at this point.
This forms an inverted U-shaped function with the optimal taste intensity mixture about 50% to
75% MSG and 25% IMP
b
IMP has little or no umami taste intensity at 100% (no MSG in the mixture)
Table 9.2 Examples of food combinations for dishes that employ the synergistic effects between
glutamate and IMP or GMP
Glutamate source IMP or GMP source Final synergistic combination
Glutamate + IMP foods
Aged cheese Tuna Tuna melt sandwich
Kombu (kelp) Katsuobushi (cured bonito) Dashi
Onions, celery, carrots Poultry Chicken soup
Dashi Pork Ramen
Aged cheese Beef Cheeseburger
Glutamate + GMP foods
Aged cheese Dried porcini Mushroom pasta
Scallop Dried morels Scallop with morel sauce
Kombu Dried shiitake Vegetarian dashi
Japanese curry, anchovies on pizza, mushroom gravy, and French onion soup. It
also explains why dashi, the base stock in Japanese cooking, is so tasty despite
being made from only two ingredients that are cooked for a short period of time
(Umami Information Center, 2016): the kombu contributes glutamate, and either the
katsuobushi contributes inosinate or, in vegetarian dashi, dried shiitake mushrooms
contribute guanylate (see Table 9.2).
9.7 Benefits of Umami in Cooking
9.7.1 Aids in Salt Reduction
Flavor-enhancing and umami-rich ingredients such as MSG offer a possible sensory

strategy to mitigate the low palatability of reduced salt. Medical evidence indicates
that reduced sodium intake in diets can improve certain disease states such as hyper-
tension and diabetes (Feldstein, 2002). Compliance with low-sodium diets is prob-
lematic due to the decrease in palatability (Roininen et al., 1996; Okiyama &
Beauchamp, 1998; Yamaguchi & Ninomiya, 2000). MSG was acknowledged in
2019 by the National Academies of Sciences, Engineering, and Medicine as a viable
strategy to reduce sodium in the food supply (Institute of Medicine, Food and
Nutrition Board et al., 2010). The amount of sodium in MSG (12.28 g/100 g) is one-
third that in salt (39.34 g/100 g), and the usage level as a food additive is quite low
(0.1–0.8% by weight) (Maluly et al., 2017; Halim et al., 2020). Adding MSG in
excess will create unbalanced flavors, decreasing palatability, so its use is self-
limiting (Yamaguchi & Takahashi, 1984). This means that MSG has the potential to
play a key role in reducing salt in food products while maintaining or increasing
likability (see Chap. 4 of this book for more detail on this topic).
9.7.2 Provides a Bass Note in Cooking
The bass note that MSG provides in cooking may not be recognizable to average
consumers, yet without it, some dishes taste like something is missing, an absence
of full flavor potential. Like the bass part in music, it may not be overtly noticeable,
but it provides the foundation that all the other parts build on—in dishes, MSG sup-
ports and enhances overall flavor. MSG has been described as “fullness of the
mouth” and “richness” (Yamaguchi & Ninomiya, 2000) (see Chap. 2 of this book
for more detail).
9.7.3 Increases Salivation
Umami, which has been shown to increase salivation (Schiffman, 1998), is notable
by persistence in the palate and salivation at the end of the palate, signs of the pres-
ence of umami. This factor has been utilized in elderly patients to increase food
intake (Sasano et al., 2015). The most important roles of saliva are during chewing
food and in maintaining oral health. Saliva helps protect the teeth and mucosa from
infection and maintain healthy taste receptors and speech communication (Uneyama
et al., 2009) (see Chap. 7 of this book for more detail).
Taste dysfunction is shown to have a negative effect on health, correlating with
poor appetite, reduced dietary intake, and weight loss (Sasano et al., 2014). This
affects the elderly population most of all (Schiffman, 2000). Taste function and sali-
vation are closely related, and the umami taste is shown to promote salivary secre-
tion. A 2015 study found that treating decreased salivation reduced hypogeusia
(Sasano et al., 2015), showing that salivation is essential to maintain normal taste
function. The increased salivary flow rate, due to the gustatory-salivary reflex from
umami taste stimulation, improved taste function, appetite, weight, and overall
health in elderly people (Sasano et al., 2014, 2015).
9.7.4 Adds “Meatiness”
While it is hard to describe what umami tastes like, it is often referred to as meaty
or savory. These flavor notes are particularly important in plant-based formulations
for creating both meatiness and depth, pleasurable sensations that are absent in
dishes that do not contain animal products (Yamaguchi, 1998a, b).
9.8 Conclusion
This chapter has focused on umami from a culinary point of view, as a basic, global
taste that has played a traditional culinary role as a source of deliciousness for mil-
lennia before it was scientifically identified. The ancient Romans added umami with
their fermented fish sauces, and the French gastronome Brillat-Savarin defined a
water-soluble substance in meat flavors, but umami compounds—glutamate, IMP,
and GMP—were not identified until 1908, by Professor Ikeda. Even so, it took
another century for scientific evidence to reveal the existence of umami receptors on
the tongue, identifying it as a bona fide taste element. These discoveries have helped
chefs and cooks around the world understand why they traditionally gravitated
toward umami-rich foods in their cuisines. Fundamental to what makes these foods
delicious is the presence of glutamate and nucleotides that, from a chemosensory
perspective, are a source of a fundamental, pleasurable taste. The analysis of umami
compounds in foods and ingredients has given light to the cooking processes and
food technologies that deliver a strong umami taste through such ingredients as
MSG. For chefs and culinary students, learning how to recognize and distinguish
the umami taste is an important part of their training in the kitchen, allowing the
mindful application of umami-rich ingredients in recipes and menus. Understanding
umami helps chefs and culinary students build balanced and complex flavor pro-
files. From a health perspective, this taste has been proven advantageous in reduced-
sodium foods, mitigating the low palatability of low-salt products, and can aid
digestion in the elderly by increasing saliva production. MSG and other concen-
trated umami ingredients can also provide the meaty and savory flavor notes that
can be especially useful in plant-based formulations. Combined, these benefits of
umami make it an attractive option to improve both health and flavor.
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Chris Koetke, CEC, CCE, HAAC, is Corporate Executive Chef at Ajinomoto Health and
Nutrition North America, Inc. He has been Executive Chef at Les Nomades in Chicago and served
for 20 years as Executive Director of the Kendall College School of Culinary Arts in Chicago and
Vice President of Culinary Arts for Laureate International Universities. He serves as the Chair of
the Feed the Planet Committee of Worldchefs and in 2010 received the inaugural Chefs
Collaborative Pathfinder Award for his work in making sustainability mainstream. He has hosted
his own national TV cooking show and coauthored the well-known culinary textbook The Culinary
Professional (2010).
Lauren Miller is a Food Scientist working for a biotechnology company focused on the sustain-
able production of a microbial-based protein source through biomass fermentation. Miller received
her BS in Culinary Arts and Food Science from Drexel University followed by a Product
Development Fellowship with Ajinomoto Health and Nutritional North America, Inc. As a
Research Assistant in the Drexel Food Lab, she worked with the Philadelphia Department of
Public Health on the Sodium Reduction in Communities Program. She contributed to research at
Monell Chemical Senses Center exploring the effects of fermentation on phytic acid and liking in
bread models of pearl millet.
Jonathan Deutsch, PhD, CHE, CRC, is Professor in the Department of Food and Hospitality
Management in the College of Nursing and Health Professions at Drexel University. He is Founding
Program Director of Drexel’s Food Innovation and Entrepreneurship programs and Vice President
of the Upcycled Food Foundation and has built the culinary arts program at Kingsborough
Community College, City University of New York (CUNY), and the PhD concentration in food
studies at the CUNY Graduate Center and School of Public Health. He directs the Drexel Food
Lab, focusing on solving real-world food system problems, and is the coauthor or coeditor of eight
books, including Culinary Improvisation (2009) and The Anti-inflammatory Family Cookbook
(2020), and numerous articles on food studies, public health, and hospitality education.
Index
A G
Appetite control, 102, 104, 117 Gut-brain, 60, 61
B H
Basic taste, 1–3, 7, 11, 14–16, 18–24, 27, 43, Health outcomes, 167, 168, 171
44, 48, 49, 51, 56, 58, 59, 62, 75, 76, Healthy diets, 165–177
80, 81, 83, 101, 103, 127, 128, 135, History of umami, 10, 44–47
148, 169, 170, 176, 184, 187, 189 Human milk, 4, 16, 17, 127, 131–138
Human perception, 3
C
Chefs, 183–185, 188–190, 193 I
CNS taste system, 62 Infant formula, 127, 131, 132, 134,
Cognitive function, 60, 148, 154–158 135, 138
Comparative taste, 11, 47, 127 Infants, 2, 12, 106, 115–116, 127, 128,
Cooking, 4, 9, 18, 77, 103, 183–185, 191–193 130–138, 172
Culinary, 183–185, 188, 189, 193
M
D Meaty, 2, 33, 86, 192, 193
Dietary guidelines, 77 Monosodium L-glutamate (MSG), 2, 3,
8–13, 20–35, 73, 80–87, 90, 102,
103, 106, 107, 111–117, 119,
F 128–130, 135, 136, 148–150, 152,
Fermentation, 4, 75, 80, 129, 183, 185, 187, 154–158, 169, 170, 172, 174, 183,
188, 190 184, 187–193
Flavor, 9–11, 14, 16, 20, 28, 33, 34, 73–76, 79, Mouthfeel, 2, 5, 62, 85, 87, 129, 174, 190
82–90, 102, 103, 106, 112, 117, 127,
128, 130, 134, 135, 138, 148–150, 152,
167, 169, 170, 174, 184–193 N
Free amino acids (FAAs), 17, 115, 116, 128, Nutrition, 5, 14, 17, 127, 132, 148, 151, 158,
129, 131–138, 170, 185–187 165–168, 177, 191
© The Editor(s) (if applicable) and The Author(s) 2023 197

https://doi.org/10.1007/978-3-031-32692-9
198 Index
O Sodium chloride (NaCl), 1, 8, 9, 11, 16,

Oral sensation, 5, 11, 28, 34, 61 20–28, 31–33, 44, 73–76, 79–81, 83,
84, 86–88, 135, 150, 156
Sodium reduction, 73, 78, 80, 84–90
P
Plant-based, 4, 165, 166, 174, 175, 177,
192, 193 T
Protein regulation, 55, 103, 116, 118 Taste, 1–5, 7–12, 14–16, 18–34, 73–76,
Psychophysics, 128–129 79–90, 101–103, 117, 120, 121,
127–130, 134–136, 138, 148–154, 157,
158, 165, 167, 169–174, 176, 184, 185,
R 187–190, 192, 193
Receptors, 3–5, 14–16, 23–26, 28, 30, Taste evolution, 170
32–34, 50–54, 57, 61, 76, 101, Taste qualities, 1–4, 8, 11, 16, 24, 27, 32, 43,
104, 119, 128, 129, 138, 150, 153, 44, 48, 57, 89, 101, 127, 138, 153, 169
154, 158, 169–174, 176, 184, 189, Taste receptors, 2, 14, 23, 28, 34, 43, 48,
190, 193 51–55, 57, 76, 101, 102, 119, 127, 128,
Recipe creation, 187, 190 169–174, 184, 189–190, 192
Transduction, 48, 49, 51, 52, 54, 55, 76, 154
S
Salivary function, 150 U
Salt, 7–9, 20, 21, 23, 73–80, 84–90, 102, 127, Umami, 1–5, 7–10, 12, 14–35, 73, 75, 80–90,
128, 148–150, 166, 168, 169, 174–177, 101–121, 127–136, 138, 148–158, 165,
184, 187, 188, 191 167–177, 183–193
Satiation, 2, 128–130, 134–138 Umami history, 184–185, 189
Satiety, 2, 34, 102–121, 128, 129, Umami taste, 3–5, 7–9, 13–16, 20–35, 43, 44,
136–138 47, 48, 50–54, 56–59, 62, 73, 80, 82,
Savory, 2, 4, 28, 80, 85, 102, 103, 83, 85, 87, 90, 101–106, 115–121,
112, 114, 116, 117, 119, 128, 127–130, 134, 138, 148, 151–154, 157,
129, 134, 135, 170, 174, 184, 187, 158, 167, 169–170, 172, 176, 184, 186,
188, 192, 193 187, 189, 190, 192, 193

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Food and Health

Ana San Gabriel

ISSN 2509-6389 ISSN 2509-6397 (electronic)

Paper in this product is recyclable.

is a strong interest in understanding factors controlling satiety and satiation. It is

Tokyo, Japan Ana San Gabriel

Gary Beauchamp Monell Chemical Senses Center, Philadelphia, PA, USA

© The Author(s), 2024 1

1.1 Species Differences in Umami Perception

Do other species detect umami? Taste qualities are human-derived percepts, so it

1.2 Umami Perception in Humans

Why, from a functional and evolutionary perspective, is human perception of umami

Beauchamp, G. K. (2009). Sensory and receptor responses to umami: an overview of pioneering

Ryusuke Yoshida and Yuzo Ninomiya

2.1 Historical Context of Umami and MSG

2.1.1 Discovery of Umami Taste by Kikunae Ikeda

© The Author(s), 2024 7

Fig. 2.1 Dr. Kikunae

(mannitol, sodium, and potassium chloride), and lead precipitation, he finally

2.1.2 First Symposium on MSG by the US Army

2.1.3 Chinese Restaurant Syndrome and MSG Safety

1908: Ikeda’s Japanese patent

1913: Finding of umami taste of 5’-inosinic acid

1920 1920s: Use of MSG in army rations by Japanese Imperial Army

1930s: Prevalence of consumption of MSG in USA

1948: First symposium on monosodium glutamate in Chicago, USA

1955: Second symposium on monosodium glutamate in Chicago, USA

1960 1960: Finding of umami taste of 5’-guanylic acid

1968: Chinese restaurant syndrome

1978: First international conference on glutamic acid in Milan, Italy

2008: Umami symposium in Tokyo

Fig. 2.2 The chronology of umami taste and monosodium L-glutamate

2.1.4 Umami as a Basic Taste

In 1978, the first international conference on glutamic acid (“The International

2.2 Umami and Other Basic Tastes

2.2.1 Umami Substances in Foods

Table 2.1 Amino acid composition of selected foods

Table 2.2 Free glutamate in selected foods

Food product Free glutamate (mg/100 g)

Table 2.3 5′-Ribonucleotides in foods

Table 2.4 Extractive components in the leg meat of snow crab

Component mg/100 g Component mg/100 g Component mg/100 g

2.2.2 Interaction Between Umami and Other Tastes

2.2.3 Psychophysical and Multidimensional Studies

This concept has been adopted to show the independence or distinctiveness of

2.2.5 Neural Pathways for Umami Taste

2.3 Differences in Umami Taste

2.3.2 Tongue Regional Differences

Sensitivity of the tongue differs by region. In experimental animals, these regional

umami substances by gurmarin (Ninomiya et al., 2000; Yasumatsu et al., 2006). In

2.4 Distinctive Phenomena of Umami Taste

2.4.1 Intensity of Umami Taste

As mentioned by Ikeda, one of its characteristics that distinguishes umami from

100 Quinine Tartaric

Tartaric acid: S = 14.45 log2(x/0.00296)

As described earlier in this chapter, characteristic descriptions of umami taste often

2.5 Conclusion and Perspective

Ryusuke Yoshida is Ph.D. He is Professor of Oral Physiology in the Faculty of Medicine,

Eugene R. Delay and Stephen D. Roper

© The Author(s), 2024 43

Tokyo, Japan Ana San Gabriel

1.1 Species Differences in Umami Perception

1.2 Umami Perception in Humans

2.1 Historical Context of Umami and MSG

2.1.1 Discovery of Umami Taste by Kikunae Ikeda

2.1.2 First Symposium on MSG by the US Army

2.1.3 Chinese Restaurant Syndrome and MSG Safety

2.1.4 Umami as a Basic Taste

2.2 Umami and Other Basic Tastes

2.2.1 Umami Substances in Foods

2.2.2 Interaction Between Umami and Other Tastes

2.2.3 Psychophysical and Multidimensional Studies

2.2.5 Neural Pathways for Umami Taste

2.3 Differences in Umami Taste

2.3.2 Tongue Regional Differences

2.4 Distinctive Phenomena of Umami Taste

2.4.1 Intensity of Umami Taste

2.5 Conclusion and Perspective

3.1 A Brief History of Umami

3.2 Umami Psychophysics: Humans and Rodents

3.3 Overview of Tongue and Gustatory System

3.5 Structure and Function of Umami Receptors

3.7 Cranial Nerve Responses to Umami

3.8 Nucleus of the Solitary Tract and Parabrachial Nucleus

3.11 Umami Signaling in the Gut: Gastrointestinal System

3.12 Summary and Conclusions

4.2 Role of Salt in Food

4.2.2 Major Roles of Salt

4.3 Sodium Intake and Health

4.3.1 State of Sodium Consumption

4.3.3 Sodium’s Effects on Health

4.4 Strategies to Reduce Sodium Intake

4.4.3.3 Umami Compounds as Flavor Enhancers

5.1 Why We Taste Umami: Lessons from the Diet

5.2 Umami and the Regulation of Protein Intake

5.2.1 Regulation of Protein Intake

5.2.2 Protein: The Most Satiating Macronutrient?

5.3 A Role for Umami in Protein-Based Satiety?

5.3.1 Umami-Enhanced Satiety: Human Experimental Studies

5.3.2 Glutamate and Satiety in Human Infants

5.3.3 Protein Need State, Satiety, and Liking for Umami Taste

5.4 Possible Models for Umami-Enhanced Satiety

5.5 Uncertainties and Future Directions

6.1 Umami Taste in the Oral Cavity and Alimentary Canal

6.1.1 Oral Cavity and Taste Psychophysics

6.1.2 Alimentary Canal and Feeding Behaviors

6.2 Inborn Responses to Umami Taste