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ORIGINAL
Dietary supplementation with monosodium glutamate withdietary balance such as protein, salt and sugar intake withincreasing T1R3 taste receptor gene expression in healthy
females
Kana Beppu†1, Hitoshi Shono†2, Ayuka Kawakami1, Tomoe Takashi1, Suzuno Watanabe1, Akari Yoshida1,
Masashi Kuroda1, Chisa Fujimoto2, Ryo Kanamura2, Hiroki Ohnishi2, Eiji Kondo2, Takahito Azuma2, Go Sato2,
Yoshiaki Kitamura2, Rie Tsutsumi1, Hiroshi Sakaue1, and Noriaki Takeda2
Department of Nutrition and Metabolism, Institute of Biomedical Sciences, Tokushima University, Tokushima, Japan, 2Department of Otolaryngology, Institute of Biomedical Sciences, Tokushima University, Tokushima, Japan
1

Abstract : We previously showed that chemotherapy-induced dysgeusia was associated with lingual taste receptor gene expression, and monosodium glutamate (MSG) improved dysgeusia by upregulating taste 1 receptor 3(T1R3) gene expression. In recent years, decreased taste sensitivity has also been reported in some young people,
and these are partly due to their disordered eating habits. From these background, we investigated the effects of
MSG supplementation on taste receptor expression and dietary intake in healthy females. Fifteen young healthy
volunteers were enrolled for the present crossover study and divided in two groups (dietary supplementation
with MSG at 2.7 g / day or 0.27 g / day). The relative expression of T1R3, a subunit of both umami and sweet taste
receptors, in the tongue was assessed by quantitative PCR analysis. Food intake was assessed by food frequency
questionnaire (FFQg), and body composition was measured using Omron HBF-701. T1R3 expression levels in the
tongue and taste sensitivity increased significantly in participants who consumed <10 g of MSG daily, whereas
no alteration was observed in participants who consumed >10 g of MSG daily. Furthermore, protein, fat, andcarbohydrate (PFC) balance and salt and sugar intake improved by MSG supplementation. In conclusion, MSG
supplementation increased T1R3 expression in the tongue and improved dietary balance. J. Med. Invest. 68 : 315320, August, 2021
Keywords : monosodium glutamate (MSG), taste 1 receptor 3 (T1R3), dysgeusia

INTRODUCTION
Dysgeusia is serious because it lowers quality of life and exacerbates pathological conditions, including weight loss and malnutrition. Taste sensitivity is an important factor for maintaining
life as well as for enjoying diet (1). In addition to medication andaging, there are several causes of decreased taste sensitivity.
Cancer therapy, diabetes, infection and inflammation, vitamin
B12 or zinc deficiencies, or dry months are well-known causes.
In addition, COVID-19 induces chemosensory loss, such as lossof taste and smell. Many cases of dysgeusia caused by COVID-19
have been reported worldwide (2-6).
Sense of taste is critical for the identification of healthy nutrients and avoiding unhealthy compounds, such as spoiled or
poisonous food. However, even in young people, taste sensitivity
is sometimes lowered, without the being person aware of it (7). In
general, reduced taste sense leads to inadequate dietary intake,resulting in malnutrition or overeating, whereas low levels oftaste sensation are also caused by food habits in healthy subjects(8).
The basic taste modalities are sweet, sour, bitter, salty, andumami, which are perceived by specific taste receptors in
the taste buds on the tongue (9). Umami taste was identified
Received for publication June 16, 2021 ; accepted June 29, 2021.

more than 100 years ago by a Japanese scientist, Dr. Kikunae
Ikeda ; however, it has been recognized as a fifth distinct taste
owing to the identification of its receptors in the last two decades(10-12). Umami and sweet taste receptors are heterodimeric proteins composed of taste 1 receptor 1 (T1R1) and T1R3 and T1R2
and T1R3 subunits, respectively (13), whereas bitter taste receptors are monomers of the 25 types of T2R subunits (14). Previously, we reported that lingual taste receptor gene expression
was associated with chemotherapy-induced dysgeusia in patientswith head and neck cancer (15). T1R3 gene expression decreases
after chemotherapy, whereas T2R5 gene expression increases,
which is correlated with decreased taste sense.
Monosodium salt of L-glutamate (MSG) is an umami compound used worldwide as a seasoning and is detected by umami
taste receptors in the tongue (16). We have recently found that
stimulation of taste buds with MSG results in upregulationof T1R1 and T1R3 expression levels in the tongue of patientswith head and neck cancer undergoing chemotherapy. Here, we
analyzed the effect of MSG supplementation in healthy young
females. MSG was added to rice at each meal three times per
day for seven days. A mucosal scraping sample was obtained
from the foliate papillae on the tongue of study subjects to assess
T1R3 abundance by real-time PCR analysis. In addition, we
examined the effects of dietary supplementation with MSG on
daily dietary intake.

†

These authors contributed equally to this work.
Address correspondence and reprint requests to Rie Tsutsumi, PhD,
Department of Nutrition and Metabolism, Institute of Biomedical
Sciences, Tokushima University, 3-18-15 Kuramoto, Tokushima 7708503, Japan and Fax : +81-88-633-7113.

The Journal of Medical Investigation Vol. 68 2021

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K. Beppu, et al. monosodium glutamate increases T1R3 expression

SUBJECTS AND METHODS
Subjects and study design

The study was approved by the Tokushima University Hospital Committee for Medical Ethics (approval number : 2405), andwritten informed consent was obtained from each patient prior
to enrollment. The Study design complied with the guidelinesof Declaration of Helsinki, and all methods were performedin accordance with the relevant guidelines and regulations of
the Institutional Ethics Committee of Tokushima University
Hospital, Japan. The study was designed as a crossover study.
Fifteen healthy young participants were enrolled in the study.
The participants were randomly divided into two groups : one
group included eight participants who were supplemented with
2.7 g / day (0.9 g / meal) MSG for the first week. After 2 weeks ofwash out, they were then supplemented with 0.27 g / day (0.09
g / meal) MSG for a week. Another group included other sevenparticipants who were supplemented with 0.27 g / day MSG for
the first week and 2.7 g / day for another week. MSG was provided in the form of seasoning, with a double-blind addition of aseasoning containing 0.9 g or 0.09 g of MSG.

Dietary MSG supplementation
The patients were randomly divided into two groups. One
group (group A) comprised seven participants (all seven were
women) and another group (group B) comprised eight participants (of which seven were women). MSG powder was added
directly to rice as a seasoning provided by Marumiya Co. Ltd.
(Tokyo, Japan) (Table 1), and the powder was sprinkled over rice.
The powder contained 0.09 g or 0.9 g of MSG for consumption
three times per day for seven days.

Table 1. Nutritional composition of seasoning (0.9 g MSG)
Nutrients

amountenergy13 kcalprotein

1.3 g

fat

0.4 g

carbohydrate

1.0 g

salt equivalent

0.5 g

sodium

200 mg

water

0.5 g

ash

0.5 g

glutamate843.6 mg

RT-PCR analysis
The surface of the foliate papillae on the tongue of patients was
scraped with a small spatula after local anesthesia with 4% lidocaine to collect a sample of the lingual mucosa (24). Scraping was
performed immediately before and one week after the first andsecond doses of chemotherapy. All tissue scrapings were immediately mixed with RNAlater solution (Ambion, Austin, TX, USA),and RNA was extracted using an RNAqueous phenol-free RNA
isolation kit (Ambion) and amplified using the CellAmp Whole
Transcriptome Amplification Kit Version 2 (Takara Bio, Shiga,
Japan). Total RNA (1 µg) was reverse-transcribed in a final volume of 20 µL using the Primescript RT Reagent Kit (Takara Bio).
The resulting cDNA (50 ng) was subjected to real-time PCR using
specific primers in a final volume of 10 µL using StepOnePlus

Real-Time PCR System (Life Technologies, Waltham, MA,
USA). The sequences of the primer sets used (forward and reverse, respectively) were 5′-TTCCCCCAGTACGTGAAGAC-3′
and 5′-CAGAGAACGTCTGGTGGTGA-3′ for human T1R3
and 5′-GA A ATCCCATCACCATCT TCCAGG-3′ and 5′GAGCCCCAGCCTTCTCCATG-3′ for human glyceraldehyde-3-phosphate dehydrogenase (GAPDH) (Invitrogen,
Waltham, MA, USA). PCR products were quantified by fit-point
analysis, and the expression of T1R3 was normalized to that of
GAPDH.

Daily dietary intake
Dietary intake was calculated by a registered dietician byassessing food frequency questionnaire (FFQg) version 4.0.
Average energy and all nutrients intake over seven days wascompared between the weeks before and after the first and second doses of supplementation. To assess glutamate intake, we
also assessment dietary intake by 24-hour diet recall assessment
and calculated amount of glutamate using Japan Food Standard
Component Table 2015, An Amino Acid Component.

Body composition
Body weight and body composition were assessed using a
TANITA dual-frequency body composition analyzer (TANITA
320).

Statistical analysis
Data are presented as mean ± standard deviation. Paired
t-test was performed after assessing the normality of data. Multiple comparisons were performed using the Kruskal–Wallis test
and two-way analysis of variance. If an overall significant difference was detected, the Tukey–Kramer test was applied to identify pairs showing significant differences. All statistical analyses
and graph generation were conducted using JMP (SAS Institute,Tokyo, Japan) or PRISM 7 software (GraphPad Software, San
Diego, CA, USA). Statistical significance was set at p < 0.05.

RESULTS

Participant characteristics
A total of 15 healthy participants [21–26-years-old (23.2 ± 1.6
years)] with no medication were enrolled in the study (Table 2).
There were no differences in body composition and energy intake
between the groups before supplementation. The abundance ofT1R3 varied from individual to individual and was not associated with nutritional intake, such as sugar and glutamic acid,and dietary intake, such as sugar, confectionery, and favorite
beverages (data not shown).

Change in food intake and body composition and T1R3 expression
after MSG supplementation
We compared energy and food intake before and after supplementation with MSG. Intake of 0.27 g / day MSG did notchange food intake, including energy and salt intake, whereas
after 2.7 g / day MSG supplementation, energy, salt and sugar
intake reduced (Table3, Figure1A-C). Intake of snacks weredecreased after 2.7 g MSG intervention, suggesting that intervention reduced these nutrients such as salt and sugar. Other
nutrients such as protein and fat were not altered (Table3). Body
composition, i.e., body fat, lean body fat, and intracellular andextracellular body fat did not change throughout the study (datanot shown). In addition, nausea, vomiting, and diarrhea did notoccur after supplementation.
Next, we analyzed lingual T1R3 gene expression before andafter MSG supplementation. Supplementation with 0.27 g / day

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The Journal of Medical Investigation Vol. 68 August 2021
Table 2. Characteristic of participantsGroup A(n = 8)Group B(n = 7)p

23.8 ± 1.5

22.9 ± 1.6

p = 0.3424

0/8

0/7

Demographics
Age (year)
Gender (male / female)
Anthropometrics
weight (kg)53.7 ± 6.4

54.1 ± 7.3

p = 0.2261

BMI (kg / m2)

21.4 ± 3.1

21.8 ± 2.8

p = 0.1457

body fat (%)

29.7 ± 7.6

28.9 ± 7.7

p = 0.3164

lean body mass (kg)36.5 ± 5.9

37.5 ± 9.2

p = 0.3919

Systolic BP

128.5 ± 10.5

124.5 ± 17.9

p = 0.3862

Diastolic BP

78.5 ± 12.8

80.5 ± 16.9

p = 0.2661

0

0

n.s

70.2 ± 10.6

71.3 ± 12.2

p = 0.1125

2064.0± ± 239.4

1917.5 ± 251.5

p = 0.2612

Blood pressures (BP, mmHg)

Hypertension ( > 140 / 90 mmHg)
Herat Rate
Dietary assessment
Energy intake (kcal / day)protein intake (g / day)

58.0 ± 6.9

60.8 ± 5.1

p = 0.3428

carbohydrate intake (g / day)

224.0 ± 56.5

265.5 ± 48.9

p = 0.2827

sugar intake (g / day)

9.5 ± 4.9

8.9 ± 3.7

p = 0.2812

salt intake (g / day)

8.5 ± 3.9

18.2 ± 4.5

p = 0.6525

Pretreatment

After 0.27 g
MSG

After 2.7 g
MSG

Energy intake (kcal / day)

2032.0 ± 246.5

2017.4± ± 257.8

1872.4 ± 210.8*protein intake (g / day)59.9 ± 6.3

61.8 ± 4.1

60.8 ± 4.1

carbohydrate intake (g / day)

223.0 ± 54.7

245.5 ± 48.9

245.5 ± 40.1

Characteristics of participants before the intervention.

Table 3. dietary intake before and after MSG investigation

Fat intake (g / day)

45.6 ± 14.2

43.0 ± 11.2

41.0 ± 11.7

sugar intake (g / day)10.5 ± 4.9

10.1 ± 3.4

8.6 ± 3.1*

salt intake (g / day)11.9 ± 3.9

10.2± ± 4.5

8.1 ± 1.8*

Glutamate (g / day)15.4 ± 3.9

16.7 ± 3.1

16.3 ± 4.1*p < 0.05 versus pretreatment

Figure 1. Change of energy, salt and sugar intake during intervention.

A) Energy intake, B) Salt intake, C) Sugar intake, were investigated before and after 0.27 g / day MSG intervention and before andafter 0.27 g / day MSG intervention. One-way ANOVA and paired t test were performed. N = 15, each.

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K. Beppu, et al. monosodium glutamate increases T1R3 expression

MSG did not change T1R3 gene expression (Figure2A). After 2.7
g / day MSG supplementation, eight participants (53%) showed
increased lingual T1R3 gene expression, but it remained unchanged in other participants (Figure2B-C).

Stratified analysis of T1R3 gene expression after MSG supplementation
Therefore, further analysis was performed to clarify whysome participants exhibited increased T1R3 gene expression
but others did not. Comparing the group in which T1R3 gene
expression increased by supplementation of 2.7 g of MSG and

the group in which there was no change, there was no significant
difference in energy intake, but the group with increased T1R3
gene expression had lower protein intake (43.2 g / day vs. 67.6
g / day ; p < 0.05), and the average intake of glutamate was alsoless than 10 g per day before intervention (9.6 g / day vs. 21.4
g / day) (Figure3A-B). In addition, the group with increased T1R3
expression had higher salt and sugar intake before intervention(Figure3C-D) before intervention. In the group with increased
T1R3 gene expression, such low protein intake and high salt
and sugar intake improved after MSG addition. That is, protein

Figure 2. Lingual T1R3 gene expression during intervention.

A) Lingual T1R3 gene expression before and after 1, 2, 3 and 4 weeks of supplementation with 0.27 g / day MSG intervention. B) Lingual T1R3
gene expression before and after 1, 2, 3 and 4 weeks of supplementation with 2.7 g / day MSG intervention. N = 15, each. C) Comparison of Lingual
T1R3 gene expression between T1R3 changed and unchanged groups. One-way ANOVA were performed. Changed ; n = 8, unchanged ; n = 7

Figure 3. Comparison of protein, glutamate, salt and sugar intake amount.

A) Protein, B) Glutamate, C) Salt, D) Sugar intake amounts were showed in two group. Changed : group that T1R3 gene expression wasincreased by 2.7 g / day MSG supplementation ; closed bar (n = 8). Unchanged : group that T1R3 gene expression was unchanged after 2.7 g / day
MSG supplementation ; opened bar (n = 7). *p < 0.05, unpaired student t-test.

The Journal of Medical Investigation Vol. 68 August 2021
intake increased after the intervention of 2.7 g MSG, salt andsugar intake decreased, and was within the generally recommended dietary intake range. There was no clear difference inenergy intake between the group with increased T1R3 gene andthe group without change in T1R3 gene after MSG intervention.
This suggests that MSG intake changed the dietary habits or
dietary components.

DISCUSSION
MSG is a sodium salt of glutamic acid (glutamate, Glu). Glutamate is a non-essential amino acid, and is the most abundantamino acid found in nature (17). However, MSG is a controversial compound regarding its use in the food industry. In thisstudy, we investigated the effect of supplementation with MSG
on dietary balance and taste receptor T1R3 gene expression
in healthy females. We focused on the effects of MSG and noton those of other umami compounds, such as 5′-ribonucleotides
guanosine monophosphate and inosine monophosphate, as ourpreliminary work showed that MSG was the only compound that
increased T1R3 expression in the tongue of mice (unpublisheddata). The human T1R1 / T1R3 heterodimer is activated by glutamate (18). We previously observed that during the first roundof chemotherapy (cisplatin only or a combination of cisplatin and5-fluorouracil) during chemoradiotherapy resulted in a decreasein T1R3 expression levels in the lingual mucosa and taste sensitivity in patients with advanced head and neck cancer. These
effects were attenuated by dietary supplementation with MSG
during the second round of chemotherapy (unpublished data).
In addition, we focused on T1R3 because chemotherapy (but notradiation therapy) reportedly reduces T1R3 expression in the
tongue of patients with head and neck cancer without altering
T1R1 or T1R2 expression (15).
MSG is recognized as the main ingredient of umami taste.
Recently, the health benefits of umami has received considerable attention. The use of umami can help reduce excessive salt
intake, which contributes to cardiovascular disease. Differencesin the salt-exposed environment at birth and preference for salty
taste may affect the sense of taste (19). In addition, Mizuta et
al. demonstrated that umami taste disorder is a predictor ofobesity and suggested that umami contributes to meal satisfaction (20). Supplementation of MSG in elderly patients promotes
gastric acid and bile acid secretion and contributes to increased
dietary intake and improved nutrient absorption rate, resulting
in improved nutritional status (21, 22). Oral administration of
MSG also increased gastrointestinal mRNA and protein levelsof T1R1 and T1R3 in piglets (23). Although the precise mechanisms of these effects remain unclear, it is likely that the interaction of MSG with T1R1 / T1R3 umami receptors in the tongueof patients with head and neck cancer activates intracellularsignals that elicit the expression of T1R3, thereby compensating
for the inhibitory effect of chemotherapy.
In this study, we aimed to determine whether the increase in
T1R3 gene expression by MSG supplementation may be dependent on glutamate intake from the original diet. In subjects withlow glutamate intake and low protein intake, the state of “glutamate undernutrition” suppressed T1R3 gene expression, which
was considered to be improved by adding 2.7 g of glutamate daily.
In these subjects, in addition to low protein intake, high salt
and sugar intake was corrected by supplementation with MSG.
There is a strong association between sweet and umami tastes,and supplementation with MSG may support reduced sugar
intake (24). Although MSG includes sodium, taste satisfaction
with MSG may also have contributed to reduced salt intake.
The present study has several limitations. First, taste319

sensation was not assessed using an accurate taste test. Only
interviews were conducted regarding dysgeusia, and it was
confirmed that none of the subjects had any subjective abnormalities. However, a more accurate taste test, such as thewhole-mouth gustatory test, is required to be performed because
sometimes even young healthy subjects present dysgeusia. Second, the number of subjects was relatively small and the study
was designed as a crossover study. Third, only women were
included this study. Gender differences are also considered to
be involved in taste and gene expression, but we have not beenable to examine gender differences in this study. Thus, further
randomized clinical trials are required to confirm our findings.
In conclusion, this study showed that dietary supplementation
with MSG improves dietary balance by increasing T1R3 taste
receptor gene expression in healthy females with an unbalanced
diet.

CONFLICT OF INTEREST
The authors have no financial conflicts of interest to disclose.

ACKNOWLEDGEMENTS
We thank Marumiya Co. for preparing the seasoning powder
containing monosodium glutamate as well as the nurses and dieticians at Tokushima University Hospital for their cooperation.
This study was supported by the Japan Society for the Promotionof Science KAKENHI grants 18K11075 (to R.T.) and 19K09871(to N. T.).

REFERENCES
1.

2.

3.

4.

5.

6.

7.

8.

de Jong N, Mulder I, de Graaf C, van Staveren WA : Impaired sensory functioning in elders : the relation with its
potential determinants and nutritional intake. J Gerontol A
Biol Sci Med Sci 54 : B324-331, 1999
Altin F, Cingi C, Uzun T, Bal C : Olfactory and gustatory
abnormalities in COVID-19 cases. Eur Arch Otorhinolaryngol 277 : 2775-2781, 2020
Meunier N, Briand L, Jacquin-Piques A, Brondel L, Penicaud
L : COVID 19-Induced Smell and Taste Impairments : Putative Impact on Physiology. Front Physiol 11 : 625110, 2020
Jain A, Pandey AK, Kaur J, Kumar L, Singh M, Das S,
Purohit S : Is there a correlation between viral load andolfactory & taste dysfunction in COVID-19 patients? Am J
Otolaryngol 42 : 102911, 2021
Lechner M, Liu J, Counsell N, Ta NH, Rocke J, Anmolsingh
R, Eynon-Lewis N, Paun S, Hopkins C, Khwaja S, Kumar
BN, Jayaraj S, Lund VJ, Philpott C : Course of symptoms
for loss of sense of smell and taste over time in one thousand
forty-one healthcare workers during the Covid-19 pandemic : Our experience. Clin Otolaryngol 46 : 451-457, 2021
Kumar L, Kahlon N, Jain A, Kaur J, Singh M, Pandey
AK : Loss of smell and taste in COVID-19 infection in adolescents. Int J Pediatr Otorhinolaryngol 142 : 110626, 2021
Yoshinaka M, Ikebe K, Uota M, Ogawa T, Okada T, Inomata
C, Takeshita H, Mihara Y, Gondo Y, Masui Y, Kamide K,
Arai Y, Takahashi R, Maeda Y : Age and sex differences inthe taste sensitivity of young adult, young-old and old-old
Japanese. Geriatr Gerontol Int 16 : 1281-1288, 2016
Nagai A, Kubota M, Katayama Y, Kojima C : Evaluationof taste acuity by the filter-paper disc in Japanese young
women : the relationship with micronutrients status. Asia

320

K. Beppu, et al. monosodium glutamate increases T1R3 expression

Pac J Clin Nutr 21 : 406-410, 2012
Li F : Taste perception : from the tongue to the testis. Mol
Hum Reprod 19 : 349-360, 2013
10. O’Mahony M, Ishii R : A comparison of English and Japanese taste languages : taste descriptive methodology,
codability and the umami taste. Br J Psychol 77 ( Pt 2) : 161174, 1986
11. Kinnamon SC, Vandenbeuch A : Receptors and transduction of umami taste stimuli. Ann N Y Acad Sci 1170 : 55-59,
2009
12. Kinnamon SC : Umami taste transduction mechanisms.
Am J Clin Nutr 90 : 753S-755S, 2009
13. Xu H, Staszewski L, Tang H, Adler E, Zoller M, Li X : Different functional roles of T1R subunits in the heteromerictaste receptors. Proc Natl Acad Sci U S A 101 : 14258-14263,
2004
14. Mueller KL, Hoon MA, Erlenbach I, Chandrashekar J,
Zuker CS, Ryba NJ : The receptors and coding logic for bitter taste. Nature 434 : 225-229, 2005
15. Tsutsumi R, Goda M, Fujimoto C, Kanno K, Nobe M,
Kitamura Y, Abe K, Kawai M, Matsumoto H, Sakai T,
Takeda N : Effects of chemotherapy on gene expression of
lingual taste receptors in patients with head and neck cancer. Laryngoscope 126 : E103-109, 201616. Shoji N, Kaneta N, Satoh-Kuriwada S, Tsuchiya M,
Hashimoto N, Uneyama H, Kawai M, Sasano T : Expression of umami-taste-related genes in the tongue : a pilot
study for genetic taste diagnosis. Oral Dis 21 : 801-806,
2015
17. Zanfirescu A, Ungurianu A, Tsatsakis AM, Nitulescu GM,
9.

Kouretas D, Veskoukis A, Tsoukalas D, Engin AB, Aschner
M, Margina D : A review of the alleged health hazards ofmonosodium glutamate. Compr Rev Food Sci Food Saf
18 : 1111-1134, 201918. Li X, Staszewski L, Xu H, Durick K, Zoller M, Adler E :Human receptors for sweet and umami taste. Proc Natl
Acad Sci U S A 99 : 4692-4696, 2002
19. Morita R, Ohta M, Umeki Y, Nanri A, Tsuchihashi T,
Hayabuchi H : Effect of Monosodium Glutamate on Saltiness and Palatability Ratings of Low-Salt Solutions inJapanese Adults According to Their Early Salt Exposure or
Salty Taste Preference. Nutrients : 13, 2021
20. Mizuta E, Kinugasa Y, Kato M, Hamada T, Yamamoto K,
Hisatome I : Umami taste disorder is a novel predictor ofobesity. Hypertens Res 44 : 595-597, 2021
21. Toyama K, Tomoe M, Inoue Y, Sanbe A, Yamamoto S : A
possible application of monosodium glutamate to nutritionalcare for elderly people. Biol Pharm Bull 31 : 1852-1854, 2008
22. Yamamoto S, Tomoe M, Toyama K, Kawai M, Uneyama
H : Can dietary supplementation of monosodium glutamate
improve the health of the elderly? Am J Clin Nutr 90 :844S-849S, 2009
23. Zhang J, Yin Y, Shu XG, Li T, Li F, Tan B, Wu Z, Wu
G : Oral administration of MSG increases expression of glutamate receptors and transporters in the gastrointestinal
tract of young piglets. Amino Acids 45 : 1169-1177, 201324. Shim J, Son HJ, Kim Y, Kim KH, Kim JT, Moon H, Kim
MJ, Misaka T, Rhyu MR : Modulation of sweet taste byumami compounds via sweet taste receptor subunit hT1R2.
PLoS One 10 : e0124030, 2015