Trends in Analytical Chemistry 127 (2020) 115876

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Trends in Analytical Chemistry

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Characterization and evaluation of umami taste: A review

Wenli Wang, Xirui Zhou, Yuan Liu*
Department of Food Science &Technology, School of Agriculture and Biology, Shanghai Jiao Tong University, Shanghai, 200240, China

a r t i c l e i n f o a b s t r a c t

Article history: Umami, as an alimentary taste, is an indicator for the presence of valuable amino acids or oligopeptides.
Available online 30 March 2020 Umami substances, including free amino acids, nucleotides, peptides, organic acids and their derivatives,
are widely distributed in foods. They usually improve the overall taste of food such as modulating sweet
Keywords: taste, enhancing salty taste and suppressing sourness and bitterness. Though umami receptors have been
Umami molecules discovered, the evaluation of umami intensity with sensitivity and selectivity is nevertheless a challenge
Food source
because of its multivariate taste characteristics in complex food matrix source. Based on the available
Taste characterization
literature, this review summarized the distribution traits in food sources, taste characteristics, evaluation
Interaction mode
Evaluation method
approaches and the related investigations. In further studies, with advance in the field of materials,
microfluidic and nano-technologies, new evaluation approaches should be designed with the advantages
of fast, sensitive and selective, and allowing more structurally versatile umami (enhancing) ingredients
would be elucidated from raw food materials.
© 2020 Elsevier B.V. All rights reserved.

1. Introduction discovered and identified in recent years, including free L-amino

The tongue is capable of sensing nutrients and toxins through
taste receptors and signaling mechanisms. Two important taste
receptors for sensing nutrients have been identified in tongue
belonging to the taste 1 receptor (T1R) family, which gives rise to
sweet and umami sensations. Members of the T1R family mainly
respond to particular taste stimuli by the formation of hetero-
dimers (T1R1/T1R3 that responds to umami and T1R2/T1R3 that
responds to sweet) [1]. Umami has with the property of enhancing
food palatability, which specifically can be described as ‘savory’ or
‘meaty’, originating from the intake of protein-rich food products
[2]. Umami not only improves the taste of a variety of foods, but also
plays a role in human metabolism [3]. It is known to enhance
appetite and satiety [4], ameliorate eating disorders [5], remediate
hypogeusia [6], and to increase the peristaltic reflex and pellet
propulsion through the distal colon [7]. Umami have been estab-
lished as a basic taste, belonging to the same category containing
sweet, sour, salty and bitter taste [8].
As a result of the multiple binding sites of ligands on the umami
heterodimer (T1R1/T1R3), umami receptor shows low specificity so
that it can respond to multiple, chemically-diverse umami mole-
cules. A series of structurally diverse umami compounds have been
* Corresponding author.
E-mail address: y_liu@sjtu.edu.cn (Y. Liu).
acids, purine nucleotides, peptides, organic acids, amides and their
derivatives [9,10]. The discovered first umami agent was mono-
sodium glutamate (MSG) that was separated from brown kelp by Dr
Kikunae Ikeda (1908) [11]. Additionally, peptides and their de-
rivatives having nutritional or bio-active properties exhibited the
umami (enhancing) sensory attribute through different modes of
interactions with the umami receptor [10,12e14]. Other umami
(modulating) compounds, such as amides and derivatives of 50 -ri-
bonucleotides (IMP or GMP), were discovered as novel umami
enhancers, and they were generated by chemical synthesis [15e17].
Moreover, their umami (enhancing) characteristics were further
compared by structure-activity relationship [9]. Further unknown
umami-active compounds were identified and purified from a
complex food matrix through sophisticated analysis and separation
techniques. These separation techniques included macro-porous
resin adsorption, membrane separation and chromatography [18].
High performance liquid chromatography-mass spectrometric
(HPLC-MS) and nuclear magnetic resonance (NMR) coupled with
human sensory evaluation provide a better approach for the
identification of substances that contribute to umami taste. For
instance, many amides and nucleotide compounds have been
successfully identified through these technologies [9].
Umami, as a basic taste, shows multivariate characterization.
Umami substances can interact with each other and also can
interact with other taste substances to have a synergistic effect or a
suppression effect on overall taste perception. The most prominent

https://doi.org/10.1016/j.trac.2020.115876
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2 W. Wang et al. / Trends in Analytical Chemistry 127 (2020) 115876
synergistic effect is the umami enhancement of the response to L-
glutamate by 5-ribonucleotides. A study showed that the mixture
of glutamate and IMP (1:1) produced 7-fold greater umami in-
tensity compared to the intensity of glutamate alone [19]. The
molecular mechanism of the synergy can be well elucidated using
the ligand-binding model involving the Venus flytrap domain of
T1R1 by molecular modeling and site-directed mutagenesis [20].
The synergistic effect between L-theanine and IMP for umami was
investigated by human sensory evaluation and gustatory nerve
recording, and the result showed that the intensity of the umami
taste (a mixture of 25 mM L-theanine and 0.5 mM IMP) increased by
57% compared with the intensity from 25 mM L-theanine alone
[21]. Additionally, Naþ and Cl are also reported to significantly
increase the overall umami intensity of morel mushroom and dried
bonito stock [22,23]. To an extent, umami (enhancing) peptides
have had a synergistic effect with salt and MSG [24,25]. Owing to
the action of chemical molecules/treatments, including the changes
of temperature or pH and the influences of salt or polysaccharides,
the umami perception usually be suppressed [18,26]. Additionally,
umami can modulate sweet taste, enhance salty taste and suppress
sourness and bitterness [27e29]. Therefore, the umami intensities
of compounds or the complex food sample cannot easy be evalu-
ated due to its multivariate taste characteristics. The known umami
molecules can be measured by the conventional methods, such as
human sensory evaluation, taste activity values (TAVs) and equiv-
alent umami concentration (EUC). Novel evaluation approaches
based on stoichiometry and bio-sensing techniques have been
developed to match with specific umami molecules. For instance,
the binding ability of umami ligand and receptor T1R1/T1R3 from
cat was quantified by intrinsic tryptophan fluorescence [30]. In
addition, a study investigated the synergistic effects between
umami peptides, monosodium glutamate and taste receptor T1R1/
T1R3 by molecular docking and simulation [24]. The taste cell from
taste buds or epithelium was employed to detect extracellular po-
tential of taste cells stimulated by umami substances, e.g. MSG, but
it also responded to the stimulation by others four tastes [31,32].
Therefore, the umami intensities could not be adequately evaluated
for the complex food model especially for those containing un-
known umami active compounds. Thus, more evaluation modes
should be developed in further studies.
A number of reviews on the umami taste have been conducted
worldwide. They mainly focused on umami (enhancing) com-
pounds and its receptor [9,10], processing methods of umami
substances [18], the criteria of umami as a basic taste [8], the effects
of umami ingredients on human health [33], and umami sensing in
gastrointestinal tract [34]. However, to date there is no detailed
summary regarding the taste characterization and evaluation ap-
proaches on umami substances. In this review, we have discussed
recent investigations on effects of umami substances on the dis-
tribution traits in food source, taste characterization and their
evaluation approaches with overall conclusions based on these
findings.
2. Umami substances in food
2.1. The main umami (enhancing) substances and their food source
Several traditional foods are rich in umami substances, such as
the fermented animal-based/plant-based products from fish, milk,
beans, grains, mushrooms, and tea. Based on the type of food
source, the main umami substances are summarized in Table 1.
Edible mushroom is a typical and peculiar umami food source. The
umami ingredients from 34 edible mushroom species were sum-
marized based on their species type, maturity stage, quality grade,
storing time, processing methods and parts of mushroom (pileus,
stripes, fruit body and mycelia) [35]. The findings showed that the
common umami ingredients of edible mushroom are MSG-like
amino acids (Asp, Glu) and 50 -nucleotides (GMP, IMP and XMP)
[35]. Aquatic products are another typical food with rich umami
components. Key taste active compounds from 24 known aquatic
products were summarized in the study of Zhang [36]. Amino
acids (Glu, Gly, Ala and Arg), 50 -nucleotides (GMP and IMP) and
inorganic ion (Naþ, Kþ, Cl and PO43) were found in most of
these aquatic products. L-theanine, which is a derivative of Glu,
was found in the green tea [37,38]. It is the major “umami” (good
taste) component of tea and has favorable physiological effects on
mammals [39].
Recently, umami oligopeptides have become a key interest in
the development of flavor-based agent on their characteristics on
umami perception. More than 100 umami (enhancing) peptides
have been identified in various foods [10] that have a strong syn-
ergy with MSG, IMP and GMP [8]. Certain peptides are also indis-
pensable elements that contribute to the specific flavors of food,
such as Takifugu obscurus [36]. We should ask what the difference
between oligopeptides and other umami molecules (MSG, GMP and
IMP) is. There is little evidence showed that they have different
binding domain/sites with umami receptor T1R1-T1R3 [88]. Also
there are different conformations of the T1R1-T1R3 hinge of venus
flytrap domains (VFD) when glutamate binds to the VFDs in the
presence of IMP/GMP or umami-enhancing peptides [24]. Further
studies are required on the special taste features of umami
peptides.
In traditional foods, amino acids, organic acids, nucleotides and
oligopeptides are all key umami molecules. In addition, novel
umami molecules were also found in vegetables [80], spice [85],
and insect [87]. The derivatives of amino acids, nucleotides and
peptides are the main novel umami molecules, such as GMP de-
rivatives [15,64,89e91], the glycoconjugates of MSG [92e95], MSG
derivatives [80], Fru-pGlu, Fru-Val, Fru-Met, pGlu-Gln and pGlu-Gly
[57,62]. Amines/amides [82,85,96] and other heterocyclic com-
pounds [16,29,97,98], can also elicit strong umami taste. Rubem-
amine from Chenopodium album and rubescenamine from
Zanthoxylum rubsecens elicit strong intrinsic umami taste in water
[82]. Though armatamide zanthosinamide, and dioxamine have not
shown any intrinsic umami taste, they still positively modulated
the umami receptor T1R1/T1R3 [82]. The hydrophobic compound
with a terminal pyridine residue and a linear structure (2-[[[2-
[(1E)-2-(1,3-benzodioxol-5-yl)ethenyl]-4-oxazolyle]methoxy]
methyl] pyridine) exhibits extremely potent umami activity
(27,000 times the umami taste of MSG). Moreover, this compound
evokes maximal taste intensity at 22.0 s by the time intensity
analysis, whereas the fact that MSG exhibits the maximum taste
intensity immediately on tasting [98].
2.2. Efficient methods of enhancing umami substance yields
Though various umami substances in foods have been isolated
and identified, their contents are often too low to meet the pressing
demands of the food industry particularly for the condiment in-
dustry. In order to obtain or produce these umami compounds
readily, certain technologies including fermentation, enzymatic/
acid hydrolysis, the microwave- or ultrasound-assisted water
extraction, Maillard reaction and chemical synthesis have been
applied [18]. Although these methods are all used for processing
umami substances from various raw materials, the related issues
also should be considered, such as its production efficiency, econ-
omy cost, time cost and environmental pollution index.
Therefore, as potential alternatives bioengineering approaches
including fermentation, enzymatic hydrolysis and biosynthesis are
used for producing umami substances. For example, the low
 W. Wang et al. / Trends in Analytical Chemistry 127 (2020) 115876 3

Table 1
The main umami compounds in various foods.

Food types Main umami compounds References

Muscle extracts/hydrolysate
Pork (longissimus) Undefined molecules [40]
Chicken soup/chicken hydrolysate Amino acids, organic acids, nucleotides, oligopeptides [41]
Pork (longissimus/biceps femoris) MSG, free amino acid, IMP, oligopeptides [42]
Beef stir-fried Oligopeptides [43]
Bovine muscle hydrolysates Undefined molecules [44]
Dried Sipunculus nudus Glycine, arginine, alanine, AMP, IMP, succinic acid, sodium, potassium, phosphate and [45]
chloride ions
Snack sausages Undefined molecules [46]
Animal by-products
Porcine plasma hydrolysates Undefined molecules [44]
MRPs form bovine bone marrow Oligopeptides [47]
Seafood/hydrolysate
Labeo rohita head hydrolysates Undefined molecules [48]
Takifugu obscurus head and bone hydrolysates Amino acid and undefined molecules [49]
Fish muscle extracts Free amino acid, IMP [50]
MRPs from Takifugu obscurus muscle Amino acid and undefined molecules [51]
Brown seaseeds Free amino acid (MSG) [52]
Yangtze Coilia ectenes Glu, Gly, AMP, IMP, GMP, Kþ, Naþ, Cl, PO43- [53]
Eriocheir sinensis Free amino acid, GMP, IMP and AMP [54]
Fermented product
Korean Soybean Paste Low molecular weight acidic peptides [55]
Cheddar and Swiss cheeses L-Glu, succinic acid, propionic acids [56]
Soy Sauce Lower molecular weight fractions, free amino acids, MSG, Fru-pGlu, Fru-Val, Fru-Met, pGlu- [57,58]
Gln, pGlu-Gly
Korean soy sauce Free amino acids, Glu-enriched oligopeptides [59]
Soy sauce Peptides, Fru-Glu [60e62]
Hams Peptides [63]
Yeast Extracts Peptide, nucleotide diastereomers (R) or (S)-N-2-(1-carboxyethyl) guanosine 50 - [64,65]
monophosphate
Salted fermented fruit known as Low molecular weight fraction including L-Glu, succinic acid and organic acid [66]
asam sunti (Averrhoa bilimbi L.)
Corn sauce Glutamyl peptides, pyroglutamic acid, L-Glu, valine, N-formyl-glutamic acid, N-acetyl- [67]
glutamine
The cooked miso Dimethyl trisulfide [68]
Mushroom
Multi-mushrooms Free amino acids, 50 -nucleotides [69e72]
Shiitake mushroom Umami peptides [12]
Morel mushrooms L-Glu, L-Asp, succinic acid, (S)-morelid [22,73]
Tea
Green tea/Tea beverages Amino acids, theanine [37,74,75]
Green tea (Matcha) L-theanine, succinic acid, gallic acid, theogallin [76,77]
Vegetables
Tomatoes L-Glu, 50 -ribonucleotide [78]
Potato (Solanum tuberosum L.) MSG, aspartate, 5 0 -nucleotides [79]
Potatoes and potato chips L-pyroglutamate, monosodium D-pyroglutamate [80]
Cereal
Rice middlings hydrolysate Amino acid and undefined molecules [81]
Plant
Chenopodium album Rubemamine [82]
Nut
Peanut hydrolysate Peptides [14,83,84]
Spice
Zanthoxylum piperitum Aromatic amides [85]
Zanthoxylum rubsecens Rubescenamine [82]
Seed
Tomato seed meal hydrolysate Amino acids [86]
Insect
Silkworm pupa hydrolysate Peptides [87]

molecular weight acidic peptides naturally produced during the
fermentation of soybeans [55]. The protein hydrolysate from
cultured Takifugu obscurus by-products (fish heads and bones) with
a higher umami and lower bitterness were prepared by enzymatic
hydrolysis [49]. The recombined umami peptide (H-Lys-Gly-Asp-
Glu-Glu-Ser-Leu-Ala-OH) was prepared by gene engineering. Such
a process would increase the yields of umami peptide and offset the
disadvantage of chemical synthesized peptides [99,100]. Gene en-
gineering, in particularly, will have broad prospects for the devel-
opment of new flavor products which will provide food products
having a stronger umami taste.
3. The multivariate characterization of umami substances
The position of the individual taste stimuli on the
concentration-intensity psychophysical curve predicts important
interactions when reporting enhancement or suppression of taste
mixtures [101]. Specific chemicals elicit umami, giving food a softer
mouthfeel, a richer intensity and taste more pleasant. It also helps
to improve the overall sensory characteristics of food. Like the other
four basic tastes, perception of umami is changed when multiple
substances stimulate the receptor instead of just one. Umami
perception can be synergistic or suppressive as affected by different
4 W. Wang et al. / Trends in Analytical Chemistry 127 (2020) 115876
umami substances, and their interactions between umami and
other taste substances leading to sensory differences.
3.1. Synergistic effect
There is a significant synergy between L-glutamic acid and 50 -
nucleotide, e.g. the mixture of glutamate and inosinic acid (1:1)
produced 7-fold greater umami intensity compared with the in-
tensity from glutamate alone [19]. Additionally, this synergistic
mechanism was also reported by the ligand-binding model
involving the Venus flytrap domain of T1R1 [20] and an allosteric
effect of nucleotides on a glutamate receptor [102]. To an extent,
umami (enhancing) peptides have also showed synergism with
MSG [24,25]. Recently, Dang et al. proved the synergistic effect
between umami peptides and MSG based on taste receptor T1R1/
T1R3 by molecular docking and simulation [24], and the synergistic
effect of peptide and MSG was also evaluated by electronic tongue
[25]. L-theanine from green tea not only elicits umami taste by
activating T1R1/T1R3-expressing cells but also shows a synergistic
response with inosine 50 -monophosphate [103], when investigated
by human sensory evaluation and by gustatory nerve recording
[21]. Alapyridaine, which is isolated from heated aqueous solutions
of hexoses and L-alanine, also revealed a significant increase in
sweetness and umami character in synthetic beef taste recombi-
nate [104].
Umami synergistic effect not only exists among umami sub-
stances, but also significantly increases with supplementation of
given compounds or chemical treatments. For example, Naþ and
Cl can significantly increase the overall umami intensity of morel
mushroom and dried bonito stock [22,23]. Glutathione has
demonstrated an enhancing effect of umami and salt tastes by
magnetic resonance imaging and sensory evaluation [105]. L-Arg is
used as a strategic tool to modify sucrose perception, enhance the
umami taste, increase NaCl saltiness and caffeine bitterness and
decrease citric acid sourness for tasters [106]. Wheat gluten protein
hydrolysate showed higher umami taste, with the extent of hy-
drolysis and total free amino acid amount of Glu and Asp especially,
when beta-cyclodextrin was employed in the enzyme reaction
[107]. Deamidation by specific enzymes or acid treatments are also
very promising techniques for increasing umami taste. For instance,
wheat gluten hydrolysates were treated with glutaminase for
180min, the umami taste score increased from 1.62 to 4.27 [108].
3.2. Suppression action of umami perception
Changes in pH, addition of compounds, or chemical treatments
can affect the perception of umami, with its intensity usually being
suppressed. The intensity of umami is weakened at pH lower than
5.5 or higher than 8.0, and its intensity is also sharply reduced by
heating for an extended period of time, as is the case of L-MSG [18].
Umami perception is inhibited by sodium lactisole, which binds to
the T1R3 subunit in vitro [109]. Clofibric acid (16 mM) significantly
reduced the perceived umami intensity elicited by 20 mM MSG,
and it also inhibited the perceived umami intensity from
MPG þ GMP [110]. Polysaccharides with molecular weights of
44.9e49.7 kDa cause the suppression of the umami taste/aftertaste
in soy sauce [26]. Methional as a familiar flavor compound is an
allosteric modulator of T1R1/T1R3. An unexpected finding was that
methional served as a positive allosteric modulator of human T1R1/
T1R3 but functioned as a negative allosteric modulator of mouse
T1R1/T1R3. However, this is because that methional could poten-
tially bind at two distinct sites in the transmembrane domain of
T1R1 and that the amino acid residues in the bottom of the allo-
steric pocket engendered the switch between two modulator
modes [111]. More synergistic/suppression effects on umami are
expected to be found and elucidated by the molecular mechanisms
underlying ligand recognition in further studies.
3.3. The interactions with other tastes
Umami substances appear to enhance the perceived sweetness
or saltiness, or other flavor characteristics such as mouthfeel and
complexity [102]. However, umami substances also inhibit the
perception of sourness or bitterness. IMP and Glu enhance the
thickness and overall taste of fish muscle extracts, and also repress
the sourness sensation elicited by these extracts [50]. Due to the
multiple receptor mechanisms for umami detection and the
complexity in taste perception models for rodents, a synergistic
mixture of monopotassium glutamate and IMP were perceived by
mice as having a sucrose-like sweet taste [112]. Additionally, the
sensitivity of sucrose to the sweet receptor hT1R2/hT1R3 was
significantly attenuated by the presence of MSG and umami pep-
tides [27]. However, umami nucleotides did not change the sensi-
tivity of sucrose, so it is necessary to make a thorough inquiry on
the intrinsic reason [27]. Due to the positive correlation between
excessive intake of sodium and the incidence of hypertension,
health awareness of people has driven demand for low-sodium
foods [113]. An investigation found that umami boosts likeability
of low-sodium foods and enhances salty taste [114].
Although controlled enzyme hydrolysis techniques are widely
applied in food industry, they can produce undesirable bitterness
tastes if the hydrolysis conditions are not effectively regulated.
Methods of reducing bitterness would be a promising research
direction. The generation of umami substances may suppress bitter
signal transduction [108]. Recent studies found that bitterness can
be suppressed by umami peptides via the human bitter taste re-
ceptor hT2R16 [115]. The key umami-active fraction in current
productions of Korean soy sauce (free amino acid and Glu-enriched
oligopeptides) can reduce human-perceived bitterness in humans.
Further studies showed that umami compounds effectively sup-
pressed the intracellular Ca2þ response induced by caffeine in the
bitter receptor hT2R43 and hT2R46 [59]. N-geranyl cyclopropyl-
carboxamide as an umami taste modulator, selectively suppressed
the salicin-induced activation of hT2R16 intracellular calcium
influx, and its suppressing efficacy was stronger than that of umami
peptides [29]. Moreover, bitter taste receptor hTAS2R16 has a
binding pocket for a wide range of umami substances, not only
umami peptides [29].
There is a seemingly unintelligible question on the relation-
ship of retronasal odor, saltiness and umami. Addition of MSG to
an aroma chicken model solution enhanced the intensity of
retronasal aroma sensation [116]. Saltiness and umami enhanced
chicken and soy sauce odor intensities, but the odors did not
enhance saltiness and umami intensities [117]. The investigation
showed that the retronasal odor of dried bonito (umami-rich)
stock improved the palatability of a salt-reduced diet due to its
characteristic aroma [118]. The after-taste intensity of saltiness
and overall taste intensity were perceived as being significantly
enhanced when salt solutions at low salt concentrations (0.68%e
0.83%) containing dried bonito aroma fraction [119]. Moreover, its
umami taste intensity was also significantly enhanced by the
addition of dried bonito aroma fraction at salt concentrations
(0.68%e1.5%) [119].
4. Evaluation of umami intensity
The aforementioned molecule types having umami (enhancing)
characteristics have been identified, but suitable methods to eval-
uate their umami intensity are complex. Though both MSG and the
umami taste intensity of green tea can be evaluated through
 W. Wang et al. / Trends in Analytical Chemistry 127 (2020) 115876
potentiometric sensors with lipid polymer membrane, it is difficult
to quantify a degree of taste or establish taste standards [38,120].
Umami perception is related to chemical transduction, and the
umami receptors are responsible for sensing umami molecules.
Although numerous studies have been performed on umami, the
qualitative analysis of umami intensity remains our main objective.
The reported evaluation methods are summarized in Table 2.
4.1. Chemical index
The known umami components include free amino acids, nu-
cleotides, organic acids, sugars and related compounds, peptides
and inorganic ions and can be detected and identified by HPLC-MS
and NMR [12,15,43,121]. Their umami intensities have been eval-
uated using two important indices, taste activity values (TAVs) and
equivalent umami concentration (EUC) [45]. EUC is generally used
to express the synergistic relationship between umami amino acids
and 50 -nucleotides [122], and it is very suitable for the evaluation of
the umami intensity of foods [54,123e125].
Sometimes EUC has been used for evaluation of the overall
umami intensity or acceptability of traditional Asian foods. For
instance, a positive correlation was found between the EUC value
and umami intensity of puffer fish (R2 ¼ 0.83), and a similar positive
correlation was also found between the EUC value and acceptability
of puffer fish (R2 ¼ 0.90) [123]. TAV is often used to evaluate the
taste impact of individual taste compounds (free amino acids, nu-
cleotides, organic bases and related compounds, sugars, organic
acids, and inorganic ions). The higher the TAV, the greater taste
contribution this compound may have [22]. It can accurately
measure the quality and quantity of umami components in foods
and minimize the potentially biasing effects of brand identity and
other information influences on consumer perception. However,
EUC and TAV are only used for the known umami molecules and
cannot be used for a complex food matrix.
4.2. Human sensory evaluation
Human sensory evaluation is the conventional method for many
taste evaluation. The classical threshold theory assumes that taste
sensations depend on the intensity and the attributes of the stim-
ulus [126]. Six different methods were compared for their
discriminating ability of umami taste intensities of monosodium
glutamate and other umami solutions. These were the two-
alternative-forced choice (2-AFC), three-alternative-forced choice
(3-AFC), quantitative descriptive analysis (QDA), the taste dilution
analysis (TDA), comparative taste dilution analysis (cTDA) and taste
recognition threshold concentrations (TRTC). For example, TDA,
cTDA and TRTC were used to evaluate the taste of a broth from beef
and vegetables [104,127]. In order to evaluate the overall perceived
umami intensity in foods, an innovative quantitative evaluation
method of umami intensity based on sensory evaluation was
established [128]. In this study, the overall umami taste intensity
values of Takifugu obscurus and Ctenopharyngodon idella were
evaluated by the Steven's equation of umami taste [128].
Table 2
Main evaluation modes of umami intensity.
Evaluation modes Methods
Chemical index EUC and TAV
Human sensory QDA, TDA, 2/3-AFC
Electronic sensor Commercial
Electronic tongue
Bio-sensor Based on taste tissue/cell/umami receptor biosensor
5
Though taste panelist is well trained and calibrated, the appli-
cation of human sensory panels in the food industry is still prob-
lematic, especially for the potential toxicity of food. Also, the
subjectivity of panelists by psychology/physiology factors can be a
significant factor [129]. For the food industry, the umami intensity
of each product should be standardized. Greater reliability and
reproducibility of analytical tools need to be achieved to establish a
reproducible protocol for accurate measurement of the umami
intensity.
4.3. Commercial sensors
Electronic taste sensing as an alternative for human sensory
evaluation is useful for the quality control and monitoring of
various samples in the food industries and has been increasingly
used for the classification and identification of similar samples
especially for liquid foods [130e132]. Because of its reliability and
repeatability, it has been widely used for the evaluation of umami
intensity in various foods. However, deficiencies remain, namely
the non-specificity for quantification of multiple components in
solution [38,72,77,133]. Additionally, the electrochemical non-
enzymic sensor has also been used for the determination of
umami substance like MSG and GMP [134e137].
4.4. Sensors based on biotic recognition elements
Bio-sensing techniques based on electrochemical substances,
are composed of biological sensing elements (recognition element),
physicochemical transducers (such as oxygen electrode, field-effect
tube, photosensitive tube, piezoelectric crystal, etc.) and signal
amplification devices. Among them, biomaterials that are capable
of producing sensitive and selective analytical signals are used as
bio-recognition elements, including microorganisms, enzymes,
antibodies, antigens, cells, tissues, nucleic acids, nerves and other
bioactive substances [138e140]. These functional bio-recognition
elements not only respond to taste stimuli, but also transmit the
chemical signals. An enzyme-based electrochemical biosensor
probe with sensitivity can be used for detection of umami sub-
stances (MSG, GMP and IMP) [141e147]. However, this sensor
would not meet the requirement for analysis of mixed umami
molecules. The main umami biosensors are summarized base on
their recognition elements, transducer and target molecules
(Table 3).
With the development of material science and the discovery of
the umami receptor cell, multiform electrochemical/bio-sensing
methods have been developed to detect the umami substances.
Taste cells/tissues are well recognized as functional biological ele-
ments for the development of biosensors, but a majority of pre-
synaptic taste cells can receive the signals that respond to more
than one taste [148]. For instances, taste cells from fungiform pa-
pillas of taste buds or epithelium was used to detect extracellular
potential of taste cells stimulated by umami substances such as
MSG, but it also responded to the stimulation by another four tastes
[31,32]. Nerves innervating taste buds including chorda tympani
Pro and cons References
Only for known molecules, [22,45]
reproducible, quality and quantity
Subjectivity and unstandardization [126e129]
Only for liquid sample, non-specificity and repeatability [130e133]
Lie on recognition elements, higher sensitive and lower specify [147e155]
6 W. Wang et al. / Trends in Analytical Chemistry 127 (2020) 115876

Table 3
Summary of main umami biosensors.

Recognition elements Transducer Target compounds References

hT1R1 Based on gold nanoparticle MSG, IMP, GMP, disodium succinate [151]
T1R1 VFT Graphene-based field effect transistor (FET) L-MSG [152]
Nanovesicles Micropatterned graphene-based FET MSG [153]
Taste receptor cells Light-addressable Glumate [31]
potentiometric sensor
Taste epithelium Microelectrode arrays chip L-Glu, L-Asp, L-MSG, L-monosodium aspartate [154]
Nerves Silver plate electrode MSG, IMP [149]
Cardiomyocytes microelectrode arrays MSG [155]
Based on enzyme Electrochemical/multi-walled carbon nanotubes/gold nanoparticle MSG/IMP/GMP [141e147]

and greater superficial petrosal also responded to umami stimuli in
a manner similar to taste receptor cells [149]. Umami molecules
bind to mGluRs receptors and G-protein-coupled dimeric receptors
(T1R1/T1R3), which are expressed in Type II cells base on the
classification of morphological characters [150]. Generally, umami
receptor taste cells only react with umami substances. Currently,
the main challenges for biosensors are the preparation of umami
receptors and the coupling of taste receptors with transducers so
that biosensors with selectivity need to be developed and devised.
Therefore, the more diversified cell-based biosensor would provide
some information for the further development of umami biosensor
tool [138,140]. The more sensitive and specify umami biosensors
are fabricated to meet the specific evaluation requirements of
umami molecules.
4.5. Other
In addition to the aforementioned evaluation approaches, more
effective evaluation means have been applied for the analysis of
umami intensity. Expressed taste receptor hT1R1/T1R3 from hu-
man source and the G protein subunit mGa15 were used to assess
the umami-enhancing activity of guanosine 50 -monophosphate
derivatives by detecting intracellular calcium levels [15]. The
assessment of umami taste was also performed by near-infrared
spectroscopy (NIRS) combined with electronic tongue [133]. The
binding ability of the umami ligand and receptor T1R1/T1R3 from
cat source was quantified by intrinsic tryptophan fluorescence, and
the result showed that the response range of L-amino acids to
umami receptor was at the micro-molar level [30].
Also, free L-amino acids, nucleotides (and their derivatives or
reaction products), and certain peptides have contributed to umami
(enhancing) activity based on their structural properties. In order to
clarify the relationship between the structure of peptides and their
special taste characteristics, a three dimensional quantitative
structure-activity relationship (3D-QSAR) model for umami pep-
tides was built based on steric field and electrostatic field to eval-
uate their taste characterization [156]. Moreover, Liu et al. further
compared the diversity characteristics of L-amino acid, nucleotide
and peptide using molecular dynamics simulations. Two different
binding effects on T1R1 affirmed that the umami peptide had
especial taste characteristics compared with other ligands [88]. The
model of umami heterodimers T1R1/T1R3 is shown in Fig. 1. With

Fig. 1. The model of umami heterodimers T1R1/T1R3 A: Non-active conformations of (T1R1-T1R3); B: Active conformations of T1R1-T1R3 combined with umami molecules
(Modified from Giulia et al. [157]); C: The constructed model of hT1R1-T1R3 VFD, in which hT1R1/T1R3 are shown in gray and blue cartoons, respectively (Reprinted with
permission from Liu et al. [88]). The ligand binding site and key structural motifs of hT1R1 are highlighted in the zoom-in image, in which two active site residues S172 and T149 are
shown in cyan sticks, and four LB2 helices, including the residues 192e205 (helix-1), 219e232 (helix-2), 256e266 (helix-3), and 277e290 (helix-4), are highlighted in cyan, pink,
lime, and magenta, respectively. VFD: Venus flytrap domain; CRD: cysteine rich domain; TM: transmembrane.
 W. Wang et al. / Trends in Analytical Chemistry 127 (2020) 115876
the development of in silico molecular modeling techniques,
structure-based computational tools were applied to investigate
the binding mode of umami molecules and receptors and to further
rationally design new umami molecules [157].
5. Conclusion and future perspective
As a pleasant and nutritional taste, umami has received much
attention. With the development of new research methods and
growing knowledge of the umami receptor (T1R1/T1R3, mGluR 1
and 4), multi-type umami agents and enhancers have been sepa-
rated and identified. Apart from the basic umami compounds
(glutamate and aspartate, succinic acid and nucleotides), many
structurally versatile umami (enhancing) compounds have been
identified in recently. Due to the multi-various taste characteris-
tics of umami, e.g. modulating sweet taste, enhancing salty taste
and suppressing sourness and bitterness, a series of umami eval-
uation approaches based on bionic technology and biotechnology
were developed, expecting the conventional methods. However,
these approaches did not appear to meet the evaluation re-
quirements of various types of umami molecules from a complex
food system.
In further studies, the research direction of umami can focus on
some key points. (i) Salt reduction in foods is an important chal-
lenge to protect population health from chronic diseases. Umami
can boost the acceptability of low-sodium foods and enhance salty
taste, so it would be beneficial to find more umami compounds
having the ability to enable the reduction of sodium levels and yet
provide a salt-enhancing effect on foods. (ii) Although many
beneficial effects of umami on health, such as improvement of food
flavor and appetite, the lower content of umami molecules in food
and the lower extraction yield restrict the development for umami
production industry. Therefore, a succession of new structurally
versatile umami (enhancing) ingredients need to be found from
raw materials or by new umami processing methods, e.g. fermen-
tation, Maillard reaction and biosynthesis. The functions and effects
of new umami ingredients need to be also investigated. (iii)
Mimicking biological senses enables an objective evaluation for
umami sensation compared with human sensory panels. There is
an increasing demand for its selectivity, specificity and detection
range and the further improvement of its reproducibility and long-
term stability. With development of 3D bio-printing, microarray
and microfluidics technology, the bio-recognition elements with
higher property would make umami bio-sensing more sensitive,
accurate and efficient, e.g. growing cells onto 3D multifunctional
platforms would become easier, and cell microarrays would further
help in high throughput screening of umami molecules, etc. (iv) The
greater numbers of binding sites of new umami (enhancing) mol-
ecules with receptors, identified by in silico molecular modeling
techniques, would help to understand the principles of umami taste
perception, enable the design of more nutritional umami molecules
and easily evaluate their umami intensities.
Acknowledgements
This work was supported by the National Natural Science
Foundation of China (Grant No. 31901813, 31972198, 31622042).
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