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Taste Perception and Food Choices

Article  in  Journal of pediatric gastroenterology and nutrition · December 2011

DOI: 10.1097/MPG.0b013e3182473308 · Source: PubMed

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 ORIGINAL ARTICLE: HEPATOLOGY AND NUTRITION

Taste Perception and Food Choices

Rossella Negri,
Mariarosaria Di Feola,
Simone Di Domenico,
M. Giuseppa Scala,


Ginevra Artesi,
Serena Valente,
Andrea Smarrazzo,
Francesca Turco,
y
Gabriella Morini, and
Luigi Greco

ABSTRACT
(pleasant) tastes, as sweet and umami (the savory taste of some
Objectives: The extent to which variation in taste perception influences
L-amino acids), elicited by energetic compounds as carbohydrates
food preferences is, to date, controversial. Bitterness in food triggers
an innate aversion that is responsible for dietary restriction in children.
and proteins, trigger attraction towards nutritive food. Bad (unplea-
We investigated the association among genetic variations in bitter receptor
sant) tastes, as bitter and sour, evolved to detect potentially lethal
TAS2R38 and food choices in healthy children in the Mediterranean area, compounds as plant secondary metabolites or microbial toxins.
to develop appropriate tools to evaluate the relation among genetic pre- Salty taste, developed to ensure ions and water homeostasis when
disposition, dietary habits, and feeding disorders. The aims of the study life moved from sea to land, can be good or bad depending on the
were to get a first baseline picture of taste sensitivity in healthy adults and concentration of sodium and on the physiological context.
their children and to explore taste sensitivity in a preliminary sample of Ancestral feeding behavior was retained by contemporary
obese children and in samples affected by functional gastrointestinal humans, although food choices do not pertain to life or death.
diseases. Nevertheless, taste is still a critical determinant of food selection,
Methods: Individuals (98 children, 87 parents, 120 adults) were recruited especially in children. As a legacy of the prehistoric age, neophobia,
from the general population in southern Italy. Bitterness sensitivity was the refuse of new unknown food, restrains the dietary habits of
assessed by means of a suprathreshold method with 6-propyl-2-thiouracil. infant babies and is related to feeding behavioral disorders.
Genomic DNA from saliva was used to genotype individuals for 3 The taste sensors are specific receptors located in the taste
polymorphisms of TAS2R38 receptor, A49P, A262 V, and V296I. Food buds of the gustatory papillae on the tongue and soft palate. Sweet,
intake was assessed by a food frequency questionnaire. umami, and bitter tastants are recognized by G-coupled protein
Results: Children’s taste sensation differed from that of adults: we receptors encoded by the TAS1R and TAS2R taste receptors gene
observed a higher frequency of supertasters among children even in the families, whereas salty and sour compounds are detected by ion
mother–child dyads with the same diplotypes. Among adults, supertaster channels. The T1R receptors are closely related and consist of the
status was related with proline-alanine-valine (taster allele) homozygous heterodimers TAS1R2/TAS1R3, which are able to recognize a wide
haplotype, whereas supertaster children were mainly heterozygous. range of sweet-tasting compounds, and TAS1R1/TAS1R3, respon-
Regarding the food choices, we found that a higher percentage of taster sible for the perception of the umami taste of L-glutamate.
children avoided bitter vegetables or greens altogether compared with The pivotal role of bitter perception to avoid accidental
taster adults. Taster status was also associated with body mass index in ingestion of potentially harmful substances is outlined by the
boys. presence in the human genome of 25 members of the T2R gene
Conclusions: Greater sensitivity to 6-propyl-2-thiouracil predicts lower family, mapping as clusters on chromosomes 7q and 5p. Human
preferences for vegetables in children, showing an appreciable effect of T2R genes display a high degree of polymorphism potentially
the genetic predisposition on food choices. None of the obese boys was a involved in the variance of individual bitterness sensitivity; how-
supertaster. ever, the functional consequences of the most of the receptor
variants on taste sensitivity is not yet known.
Key Words: bitter taste, food choices, 6-propyl-2-thiouracil, TAS2R38 Allelic variations affecting the perception of some bitter
compounds, such as salicine, quinine, and aloin, have been reported
(JPGN 2012;54: 624–629) in the TAS2R16, TAS2R19, and TAS2R43/44 genes, respectively, but
the linkage between the genotype and the trait variance is poor (1).
TAS2R38, a receptor for the thiourea compounds phenyltiocarba-

S ense of taste evolved to discriminate beneficial foods from

environmental poisons as a critical system to ensure human
survival. This goal was achieved driving innate feeding behavior to
mide (PTC) and 6-n-propylthiouracil (PROP), is the only bitter
receptor able to explain most of the variance in human bitter taste.
On the basis of sensitivity to thiourea, the human population can be
accept nutritive food or refuse potential toxic substances. Good phenotypically classified into 3 categories: insensitive, sensitive,
and hypersensitive to bitterness. The variance of this distribution is
Received July 14, 2011; accepted November 30, 2011. explained by the haplotypes generated by 3 polymorphisms in the
From the
Department of Pediatrics, University of Naples ‘‘Federico II,’’ TAS2R38 gene accounting for 55% to 85% of the variance in PTC
Naples, Italy, and the yUniversity of Gastronomic Sciences, Pollenzo- sensitivity (2).
Bra (CN), Italy. Three nonsynonymous substitutions in the TAS2R38 gene give
Address correspondence and reprint requests to Rossella Negri, PhD,
rise to amino acid changes to the protein at residues A49P, A262V,
European Laboratory for Food Induced Diseases, University of Naples,
Naples, Italy (e-mail: [email protected]). and V296I, putatively involved in G-protein interaction and receptor
The authors report no conflicts of interest. activation. The combination of the 3 single nucleotide polymorph-
Copyright # 2012 by European Society for Pediatric Gastroenterology, isms results in 5 observed haplotypes with different worldwide
Hepatology, and Nutrition and North American Society for Pediatric distribution. In Europeans the taster haplotype proline-alanine-valine
Gastroenterology, Hepatology, and Nutrition (PAV) and the nontaster haplotype alanine-valine-isoleucine (AVI)
DOI: 10.1097/MPG.0b013e3182473308 make up the vast majority of haplotypes present; 3 additional variants

624 JPGN  Volume 54, Number 5, May 2012

Copyright 2012 by ESPGHAN and NASPGHAN. Unauthorized reproduction of this article is prohibited.
 JPGN  Volume 54, Number 5, May 2012 Taste Perception and Food Choices

as the less common alanine-alanine-valine (AAV) and the rare Their age (45F/53M) ranged from 3 to 19 years. One hundred
haplotypes proline-valine-isoleucine (PVI) and alanine-alanine- twenty healthy young adults (medical or nursing students) were
isoleucine (AAI) can be observed (2). recruited on a voluntary base, after dissemination seminars at the
In vivo and in vitro studies reported PAV as the major university. Overall, the study sample mirrored the genetic and
determinant of taster status and AVI as the major nontaster hap- cultural environment of the local urban setting. Subjects were
lotype, so individuals with 2 copies of the AVI allele are basically considered eligible for the study if they were not on dietary
nontasters, whereas individuals with 1 or 2 copies of the PAV allele restrictions, with the exception of people with celiac disease,
are medium tasters or supertasters. PROP sensitivity of heterozy- who were on a gluten-free diet. The study was scrutinized and
gous common haplotypes PAV or AVI in combination with less of a approved by the ethics committee of the University ‘‘Federico II’’
common or a rare variant appears to lie in between (3,4). of Naples. Informed consent was obtained from individuals or
The genotype-phenotype relation is not stringent; this is guardians.
partly the result of the subjective methods of identification of
the phenotype but also of the likely complexity of the genetic
factors controlling taste. More genes were suggested to cooperate in Genotyping
the control of the phenotype (1,5,6). Several reports showed that the
Genomic DNA was obtained from saliva with the phenol-
perception of bitter taste is related not only to the specific taste of
chloroform extraction method following a protocol developed in our
bitter compounds but also to the wide behavior spectrum of the
laboratory. The allelic variations of the TAS2R38 gene C145G
individual in relation to food choices (7). Hypersensitive indivi-
(rs713598), C785T (rs1726866), and G886A (rs10246939) were
duals have a more restricted diet, compared with sensitive or
genotyped by means of real-time polymerase chain reaction with
insensitive individuals (8).
7900HT fast (Applied Biosystems, Carlsbad, CA) using allele-
These issues are of paramount importance in children after
specific probes and primers (Applied Biosystems).
weaning: this is the time when many children gradually develop
To estimate the population prevalence of the different hap-
erratic food acceptance, often generating anxiety in mothers.
lotypes, we excluded from the analysis the children of the mother–
Maternal expectations may not coincide with the transmission of
child dyads because their data are not independent of those of the
only 50% of the genetic predisposition to food choices, because the
mothers and would overrepresent maternal haplotypes. The
other partner is responsible for the other 50%. The genetic predis-
expected error of genotyping was estimated to be <1% by previous
position of the child is naı̈ve, because transformations brought by
work in our laboratory.
experience, social behavior, and traditions have no time to modify
genetic predisposition. There is indeed scope to explore mother–
child dyads to verify the concordance between them and the PROP Tasting
conditioning of the genetic predisposition.
There are few studies (9) analyzing the relation among food A 2-step approach was adopted to estimate the tasting
choices, sensitivity to bitter taste, and genetics in the Mediterranean phenotype of the subjects. All of the subjects underwent a supra-
population. Such studies may help us to understand the preference threshold test to assess the sensitivity of perception at suprathres-
for bitter vegetables and other greens in this population. hold concentrations. For a random sample of the subjects, after the
The specific research question at the base of the present study first suprathreshold test, we applied the threshold method, which
is to explore the phenotype-genotype correlation in bitter taste assesses the individual’s ability to discriminate low concentrations
sensitivity in either normal adults or mother–child dyads. A further of the stimulus. Suprathreshold intensity ratings are often used
task is to explore bitter taste sensitivity in obese children and those to correctly separate medium tasters from supertasters because
affected by a variety of food-related symptoms or diseases. their distribution overlaps substantially at threshold. The detection
threshold, based on the lowest concentration at which a person
METHODS detects the presence of a sensation, was instead described as the best
method to classify individuals as tasters or nontasters.
Population The test was offered to children and adults, who fasted for at
least 1 hour. The children were instructed not to swallow the
1. A total of 120 healthy adult volunteers (medical students) þ 44 solutions and, if required, underwent brief training in the ‘‘sip
healthy children and spit’’ method. Subjects tasted in ascending order 2 solutions of
2. A total of 54 children affected by a variety of functional 280 and 560 mmol/L PROP (6-propyl-2-thiouracil; Aldrich
disturbances Chemical, Milwaukee, WI) in distilled water, rinsing with water
3. A total of 92 mother–child dyads, including 41 of the 44 before and after each solution. The reported feeling or, in extremely
healthy and 51 of the 54 affected children young children, the facial expression was recorded. The sensitivity
evaluation was performed by constructing a 4-point scale in which
Inside the second group, there were children affected by: the labels ‘‘no taste,’’ ‘‘weakly unpleasant’’ (bitter, barely percep-
Feeding disturbances (6) tible), ‘‘unpleasant’’ (bitter), and ‘‘very unpleasant or terrible’’
1.
(extremely bitter) corresponded to values between 0 and 3. Facial
2. Functional gastrointestinal disturbances (10)
reactions were referred to a 4-point hedonic scale corresponding to
3. Celiac disease (18) ‘‘no taste/neutral,’’ ‘‘weakly unpleasant/depression of mouth cor-
4. Organic diseases (familial hypercholesterolemia, diabetes, ners,’’ ‘‘unpleasant/frown and depression of mouth corners,’’ and
obesity, metabolic dysfunctions) (10) ‘‘very unpleasant/frown and grimace.’’ According to the score
5. Food allergy (10) achieved, subjects were classified as nontasters (NT, score 0–2),
medium tasters (MT, score 3–4), and supertasters (ST, score 5–6).
Forty-four unselected healthy children, with their mothers, For quality control, to estimate the intraindividual variability
were recruited consecutively at a well baby clinic in the field; of the suprathreshold test, we repeated the same test in 48 of the 144
54 children affected by a variety of disturbances were recruited healthy individuals. The intraindividual concordance between the
from outpatients in the Department of Pediatrics. first and the second suprathreshold tests was 78.7%. In addition, we

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 Negri et al JPGN  Volume 54, Number 5, May 2012

applied the detection threshold method to a random sample of 10% TABLE 1. Phenotypic classification of the sample by PROP
(n ¼ 30) of the study population. PROP threshold was determined status
using a ‘‘forced choice’’ procedure, in which individuals tasted
2 samples, 1 of water and 1 of PROP solution, and were asked to Frequency %
identify the sample with the stronger taste. Six PROP solutions,
from 0.032 to 3.2 mmol/L in distilled water, were used. Threshold Adults Nontasters 47 22.5
values were identified as the first concentration correctly selected in Medium tasters 128 61.2
2 subsequent presentations. Subjects were classified as nontasters if Supertasters 34 16.3
the threshold was 0.15 mmol/L and tasters if 0.1 mmol/L. Total 209 100.0
Through the classification of individuals by the 2 methods a Children Nontasters 22 22.9
good agreement was obtained in separating insensitive from sensitive
Medium tasters 45 46.9
subjects, because the threshold of the subjects classified as nontasters
by the suprathreshold test was 0.56 mmol/L and that of the medium Supertasters 29 30.2
tasters or supertasters ranged from 0.032 to 0.1 mmol/L. Total 96 100.0
Total 305
Food Preferences PROP ¼ 6-propyl-2-thiouracil.

The data on dietary habits were collected using a food

frequency questionnaire by the 3-day recall method. The ques-
tionnaire enquired about the consumption of vegetables, legumes, of the total diplotypes, followed by the homozygous PAV/PAV and
fruits, and sweet and savory snacks. Consumption of vegetables was AVI/AVI, accounting for 24.5% and 16.5%, respectively. Combi-
detailed to bitter vegetables such as cabbage, broccoli, asparagus, nations of a common (PAV or AVI) and less of a common or rare
and spinach and nonbitter vegetables such as artichoke, eggplant, variant represented 9%. Children were evaluated separately: the
zucchini, and legumes. Mothers filled out the questionnaire on their distribution of their haplotypes was not different from that of adults
own eating habits and those of their children. (Table 2).

Data Analyses Genotype-Phenotype Association

Differences between groups were estimated by the x2 test for To assess how genetics influences the variance of bitterness
k-independent samples, with first-degree error at 0.05. The mother– perception, we evaluated the association between the TAS2R38
child concordance was evaluated by kappa statistics stratifying by diplotypes and the intensity rating of PROP suprathreshold
taster phenotype and TAS2R38 diplotype. The relation between food solutions. A few cases with rare haplotypes were not considered
choices and bitter sensitivity was estimated crossing subjects by because of the small sample size. Table 3 shows the relation
PROP status, diplotypes, sex, or age. Differences in mother–child between phenotype and genotype. It may be noted that 85% of
dyads were also evaluated. Data were analyzed using SPSS 15.0 the individuals who carry the AVI haplotype are nontasters,
(SPSS Inc, Chicago, IL). whereas the PAV haplotype was more frequent in medium tasters
and supertasters.
RESULTS The agreement between haplotype and taster phenotypes is
94.8% overall; only in 13 of 250 (5.2%) individuals the phenotype
PROP Taster Status did not overlap with the genotype.
Three hundred five subjects underwent the phenotypic classi- The PROP concentration-intensity rating does not segregate
fication of bitter sensation by the suprathreshold method: 69 by diplotype because the medium tasters or supertasters may be
(22.6%) were nontasters, 173 (56.7%) were medium tasters, and PAV homozygous as well as heterozygous. Among the individuals
130 (20.7%) were supertasters (Table 1). carrying the PAV haplotype, the presence of 2 copies was associ-
Children’s bitter sensation differed from that of adults, as ated with the supertaster status in a similar proportion for adults and
expected. Although the frequency of nontasters was similar in the children (27.3 vs 36.4), but PAV heterozygosity was associated
2 age groups, the frequency of supertasters was nearly 2-fold in with a double percentage of supertasters in children (39.5%) versus
children (16.3% vs 30.2%, x2 ¼ 8.59; P < 0.014). There was no sex
difference in the distribution of the taste phenotypes either in adults
or children. TABLE 2. Distribution of TAS2R38 diplotypes in unrelated
individuals
TAS2R38 Genotype
Diplotype Frequency %
Two hundred fifty-three individuals were genotyped for the
variant loci TAS2R38 P49A, A262V, and V296I. To avoid bias AVI/AVI 31 16.5
caused by the presence of blood relatives in the sample, the children PAV/AVI 94 50.0
of the mother–child dyads were excluded. Haplotype analysis PAV/PAV 46 24.5
performed in 188 nonrelated individuals revealed that the common PAV/AAV 6 3.2
variants PAV and AVI made up the vast majority of alleles present AVI/AAV 8 4.3
in our population, accounting for 51% and 44%, respectively, thus Others 3 1.5
composing 95% of the sample. The less common AAV haplotype Total 188 100.0
and the rare variants AAI and AVV made up the remaining 5%.
From the combination of the 5 haplotypes, 8 diplotypes were AAV ¼ alanine-alanine-valine; AVI ¼ alanine-valine-isoleucine; PAV ¼
observed: the most frequent was PAV/AVI, which represented half proline-alanine-valine.

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 JPGN  Volume 54, Number 5, May 2012 Taste Perception and Food Choices

TABLE 3. Association between TAS2R38 diplotypes and PROP sensitivity phenotypes

Nontaster Medium taster Supertaster Total

AVI/AVI or AAV (%) 39 (84.8) 5 (10.9) 2 (4.3) 46 (100)

PAV/PAV or AVI or AAV (%) 6 (2.9) 147 (72.1) 51 (25.0) 204 (100)
Total (%) 45 (18) 152 (60.8) 53 (21.2) 250 (100)
x2 ¼ 171.0, P < 0.00. PROP ¼ 6-propyl-2-thiouracil.

adults (17.1%) with the same genotype (x2 ¼ 8.83, P < 0.002) addition, among the nontaster adults a sex difference was observed:
(Table 4). Individuals who were AVI homozygous were, as 22 of 31 (71%) female nontasters ate no legumes compared with
expected, more often nontasters in adults (26/31 ¼ 84%) and in only 3 of 15 (20%) male nontasters (x2 ¼ 10.584; P < 0.001).
children (6/7 ¼ 86%). The evaluation of bitter vegetable intakes by genotype
Concerning the subjects with an AAV haplotype, the hetero- revealed that there were no differences between AVI/AVI or
zygous AAV/AVI were similar to nontaster AVI homozygous, AVI/AAV individuals and PAV homozygous or heterozygous
rating the PROP solutions as ‘‘weak’’ or ‘‘barely detectable’’ (8 among adults, whereas 48 of the 63 children with the PAV allele
of 9 individuals), whereas 6 of 10 individuals who carried the (76.2%) consumed no bitter vegetables compared with 4 of
AAV/PAV diplotype were medium tasters and the remaining 3 7 children with nontaster genotype. Finally, the PROP status did
were supertasters, as observed for the PAV/AVI heterozygous. not influence the overall intake of fruits or salty snacks.
The genotype-phenotype analysis in the mother–child dyads With regard to the relation between the bitter sensitivity
showed that there is a high degree of concordance between mother phenotype and body mass index, in our sample of children we had
and child genotype (55/62 ¼ 88.7%) (Table 5). Similarly, the 42 normal weight, 35 overweight, and 11 obese children; never-
correlation of phenotypes between mother and child was strong theless, we observed that none of the obese boys or girls was a
(65/81 ¼ 80.3%) (Table 5). supertaster, as compared with 32% of children of normal weight
(x2 ¼ 4.88; P ¼ 0.027).

Bitter Sensitivity and Food Choices

Bitterness Perception in Children Affected by
We studied the relation between the sensitivity to bitter Symptoms or Disease
stimuli and food acceptance in the context of the dietary habits
of the Mediterranean culture. Although with uncertain concordance Although the study design had no power to explore taste
among studies, greater sensitivity to PROP predicts lower prefer- sensation in several groups of children affected by a variety of
ences for cruciferous and other green vegetables. complaints, we observed that the 6 children with feeding disorders
In our sample the overall consumption of vegetables was showed a PROP sensitivity identical to that of normal children,
low; however, we observed a meaningful difference in the fre- whereas among children affected by functional gastrointestinal
quency of bitter vegetable consumption between taster groups disorders, 33.3% were supertasters, compared with 19.2% of nor-
(Table 6) and between adults and children. The status of taster mal children. Interestingly, celiac children also had a higher
or supertaster limited the consumption of bitter vegetables more (31.6%) frequency of supertasters. Unfortunately, because of the
often in children than in adults. Of the 74 children taster or super- small sample size we could not reach statistical significance (by
taster, only 19 (25.6%) ate vegetables versus 57 of 119 (47.9%) simulation we observed that the difference could have been sig-
taster or supertaster adults (x2 ¼ 6.61; P < 0.01). nificant with a larger sample size).
Moreover, whereas among the adult males the intake of This was a pilot exploratory phase, aimed to give a prelimi-
bitter vegetables appeared loosely associated with PROP status nary look at those patients who will participate in a specifically
(10/17 ¼ 58% supertasters ate no vegetables), among adult females tailored study.
it was significantly related to the supertaster status, because 32 of
the 41 supertasters (78.0%) took no servings of bitter vegetables in
3 days (x2 ¼ 5.755; P < 0.016). DISCUSSION
No differences were observed in the children regarding the Taste sensitivity is significantly different in different popu-
intake of legumes among tasters or nontasters, whereas in the adult lations because it is related to cultural attitudes and traditions. The
sample, supertasters eating legumes were, unexpectedly, nearly present work is the first study of bitter sensitivity in a pediatric
2-fold the percentage of nontasters (x2 ¼ 3.63; P < 0.07). In population in the Mediterranean area, namely in southern Italy. Our

TABLE 4. Genotype-phenotype correlation in PAV homozygous or heterozygous individuals

Medium taster Supertaster Total

PAV/PAV (%) Adults 32 (72.7) 12 (27.3) 44 (100)

Children 14 (63.6) 8 (36.4) 22 (100)
PAV/AVI or /AAV (%) Adults 78 (82.9) 16 (17.1) 94 (100)
Children 23 (60.5) 15 (39.5) 38 (100)
Total 147 51 198
x2 ¼ 8.83, P < 0.002.

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 Negri et al JPGN  Volume 54, Number 5, May 2012

TABLE 5. Analysis of concordance between genotypes and phenotypes in the mother–child dyads

Child


Mother AVI PAVy Total

Genotype
AVI
7 5 12
PAVy 2 48 50
Totalz 9 53 62
Nontaster Taster Total
Phenotype
Nontaster 5 9 14
Taster 7 60 67
Total§ 12 69 81


AVI: AVI/AVI, AVI/AAV.
y
PAV: PAV/PAV, PAV/AVI, PAV/AAV.
z
k ¼ 0.6, 95% confidence interval 0.35 < 0.6 > 0.84.
§
k ¼ 0.268, 95% confidence interval 0.05 < 0.268 > 0.45.

aim was to observe the relation between taste sensitivity and food Nevertheless, the distribution of the taster groups in our
choices and to evaluate the weight of genetic predisposition by sample was in agreement with the frequencies estimated in the
comparing phenotypes and genotypes in the population, as well as European population (2). In addition, we observed more super-
in mother–child dyads. tasters among children than in adults, suggesting that innate bitter
Increased sensitivity to bitterness is associated with child sensitivity may be modified during life, in agreement with other
neophobia, responsible for children’s rejection of new, unfamiliar reports about the decline of PROP/PTC sensitivity in adult life, with
foods, which results in a diet particularly poor in food variety (10). increasing age (12).
The data obtained in this observational study provide a solid Sex differences in taste sensitivity were reported among
basis to explore feeding disorders and food-related diseases adults by several authors seemingly resulting from anatomical
in children. differences, in that women have more fungiform papillae and
Taste receptors and effectors that mediate gustatory signals more taste buds than men (13). Sex and age also can influence
in the oral cavity have been found in gastrointestinal mucosa, the expression of phenotype in children: it has been shown that the
suggesting a role in chemosensing that triggers the physiological percentage of nontasters does not differ by sex in young children
responses to luminal content, such as absorption of beneficial or (14,15), but at puberty more nontaster individuals are boys and
rejection of toxic foods. In this regard it is likely that genetic more taster individuals are girls (16). We did not find sex diversity
variations in taste receptors could modify food intake, leading to between taster groups, neither among preschool children nor
aberrant conditions such as feeding disorders up to obesity and among adolescents.
related metabolic dysfunctions. Because polymorphisms at the TAS2R38 locus explain the
The distribution of PROP phenotypic groups in our majority of the phenotypic variation in PROP sensitivity (17), we
population was suggestive of a model of Mendelian inheritance assessed how heritable variability in bitter taste perception predicts
of bitterness sensitivity, the nontaster individuals being almost one- the phenotypic groups in our population. Haplotype analysis of the 3
quarter of all (22.6%), and the supertasters (20.7%) and the medium TAS2R38 polymorphisms allowed us to identify 2 major forms, the
tasters roughly 50% (56.7%). taster allele PAV and nontaster allele AVI, accounting for the 95%
The mode of inheritance of PROP/PTC sensitivity remains of the sample analyzed, and the rare variants AAV, AAI, and AVV.
uncertain, showing features between a simple and a complex trait The occurrence of the TAS2R38 genotypes was similar to that of the
(11). Phenotypic variation in taste perception depends on various sample (representative of the population) of Italian ancestry, geno-
factors controlling a more general taste ability, such as sex and typed by Sacerdote et al (18).
ethnicity. The agreement between haplotype and phenotype was extre-
mely high; nevertheless, the genotype-phenotype association
TABLE 6. Bitter vegetables intake by taster groups estimated among supertasters was different in adults compared with children.
by a 3-day dietary recall Whereas the hypersensitive children were homozygous as well as
heterozygous, the hypersensitive adults were mainly homozygous.
Consumption It may be argued that the prevalent association between strongest
sensitivity to PROP and the PAV homozygous diplotype in the
Status Yes No Total adults may be related to a decrease in PROP sensitivity across adult
life (12).
Nontaster (%) 18 (31.6) 39 (68.4) 57 (100) A genetic factor able to influence the sensitivity to PROP has
Medium taster (%) 60 (44.4) 75 (55.6) 135 (100) been identified in the gustin gene, the salivary isoform of carbonic
Supertaster (%) 16 (27.6) 42 (72.4) 58 (100) anhydrase, with a suggested role in taste bud development (19).
Total (%) 94 (37.6) 156 (62.4) 250 (100) Individuals homozygous for the gustin polymorphism associated
with a fully functional protein were more frequent supertasters,
x2 ¼ 8.779; P < 0.067. whether they were PAV homozygous or heterozygous, whereas

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 JPGN  Volume 54, Number 5, May 2012 Taste Perception and Food Choices

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9. Tepper BJ, Koelliker Y, Zhao L, et al. Variation in the bitter-taste
PROP sensitivity in eating behavior, food selection appears linked receptor gene TAS2R38, and adiposity in a genetically isolated popula-
to these differences. Food preference studies showed that super- tion in southern Italy. Obesity 2008;16:2289–95.
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and express lower preference for sweet foods, sweet drinks, and ences on children’s food neophobia. Am J Clin Nutr 2007;86:428–33.
salad dressings (21). PROP tasters are also more sensitive to food 11. Kim UK, Drayna D. Genetics of individual differences in bitter taste
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tasters consume fewer vegetables and added fats than do nontasters 12. Whissell-Buechy D. Effects of age and sex on taste sensitivity to phenyl-
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In the present study vegetable consumption was overall less; 1990;15:39–57.
nevertheless, children consumed fewer bitter vegetables and veg- 13. Bartoshuk LM, Duffy VB, Miller IJ. PTC/PROP tasting: anatomy,
etables at all than adults, irrespective of age or health status. We psychophysics, and sex effects. Physiol Behav 1994;56:1165–71.
speculated that in a food culture in which the overall acceptance of 14. Keller KL, Steinmann L, Nurse RJ, et al. Genetic taste sensitivity to
6-n-propylthiouracil influences food preference and reported intake in
vegetables is low, the influence of PROP status on vegetable liking
preschool children. Appetite 2002;38:3–12.
may be weak; however, sex differences were observed between 15. Mennella JA, Pepino MY, Duke FF, et al. Age modifies the genotype-
PROP sensitivity and vegetable intake among the adults and phenotype relationship for the bitter receptor TAS2R38. BMC Genet
the children. 2010;11:60.
Tepper et al (23) showed that although PROP status influ- 16. Goldstein G, Daun H, Tepper B. Influence of PROP taster status and
ences the perception of chemically diverse bitter substances, there maternal variables on energy intake and body weight of pre-adolescents.
were no differences in liking the foods containing the same bitter Physiol Behav 2007;90:809–17.
compounds that were tested. This finding suggests that the relation 17. Wooding S. Natural selection and molecular evolution in PTC, a bitter
between the ability to taste PROP and the perception and liking of taste receptor gene. Am J Hum Genet 2004;74:637–46.
bitter and other strong-tasting foods is complex and until now not 18. Sacerdote C, Guarrera S, Smith GD, et al. Lactase persistence and bitter
completely understood. taste response: instrumental variables and Mendelian randomization in
None of the obese children was a supertaster: It is clear that epidemiologic studies of dietary factors and cancer risk. Am J Epidemiol
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obese individuals have a diminished sensitivity to bitter substances.
19. Padiglia A, Zonza A, Atzori E, et al. Sensitivity to 6-n-propylthiouracil
Children affected by functional gastrointestinal disorders and chil- is associated with gustin (carbonic anhydrase VI) gene polymorphism,
dren affected by gluten intolerance showed a higher proportion of salivary zinc, and body mass index in humans. Am J Clin Nutr
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confirmation through properly designed cross-sectional studies. 20. Yackinous CA, Guinard JX. Relation between PROP taster status and fat
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CONCLUSIONS 21. Dinehart ME, Hayes JE, Bartoshuk LM, et al. Bitter taste markers
explain variability in vegetable sweetness, bitterness, and intake.
We obtained a first complex picture of bitter taste sensitivity
Physiol Behav 2006;87:304–13.
in a population, with a strong relation between phenotype and 22. Bell KI, Tepper BJ. Short-term vegetable intake by young children
genotype. Food preferences were also linked to the sensitivity classified by 6-n-propylthoiuracil bitter-taste phenotype. Am J Clin Nutr
status, although not by a simple linear model. Our preliminary 2006;84:245–51.
data support the opportunity to explore taste status in children 23. Tepper BJ, White EA, Koelliker Y, et al. Genetic variation in taste
affected by functional gastrointestinal disorders or food-induced sensitivity to 6-n-propylthiouracil and its relationship to taste perception
diseases. and food selection. Ann N Y Acad Sci 2009;1170:126–39.

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