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Sensory Science Centre, Department of Food and Nutritional Sciences, University of Reading, Whiteknights, Reading RG6 6AP, UK
Hugh Sinclair Human Nutrition Unit, Department of Food and Nutritional Sciences, University of Reading, Whiteknights, Reading RG6 6AP, UK
a r t i c l e
i n f o
Article history:
Received 18 May 2016
Received in revised form 16 August 2016
Accepted 22 August 2016
Available online 24 August 2016
Keywords:
CD36
CA6
PROP taster status
Ice cream
Fat intake
a b s t r a c t
Emerging evidence suggests fat can be perceived as a taste. G-protein coupled receptors as well as CD36, a
fatty acid translocase, have been proposed to be involved in fat perception. Therefore, differences in number of receptors and genotype of CD36 have both been proposed to influence inter-individual fat taste
perception. Fungiform papillae density (FPD) and PROP taster status are phenotypes related to receptor
number. Previous authors have proposed an association between such phenotypes and CA6 (gustin) genotype, because the latter influences receptor cell proliferation. The effect of these factors on fat perception,
preference and intake, requires further investigation. Therefore, the main aim of this study was to investigate the effects of taste sensitivity, including both genotypic and phenotypic variation, on liking of ice
cream and dietary fat intake. Participants (n = 136) age 1855 years were recruited in the UK. Hedonic
liking results demonstrated that liking for ice cream was significantly affected by the fat content of the
sample, and by demographic factors (gender, ethnicity, age) but not by the consumers CD36 rs1761667
or CA6 rs2274333 genotype, PROP taster status nor FPD. However, categorising taste sensitivity from participant responses to salt taste alone (rather than to salt and PROP) found significant differences, with low
salt perceivers liking the high fat (20%) ice cream substantially more than medium- and high salt perceivers. This indicated that increased taste sensitivity reduced liking of high fat. Cluster analysis highlighted that one group of consumers (18%) liked higher fat ice cream, whereas another (30%) liked
lower fat ice cream compared to the 52% of consumers that liked ice cream regardless of fat content.
There was a significant association between these groups and salt taste sensitivity. Concerning recorded
dietary intake, the high-fat liker group were found to have substantially higher dairy product consumption compared to high-fat dislikers. Fat intake as a percentage of total energy intake was significantly
related to CA6 genotype, however the minor allele frequency at rs2274333 is too low to draw firm conclusions within this study population.
2016 Elsevier Ltd. All rights reserved.
1. Introduction
Overconsumption of energy dense foods, particularly high fat
foods, and specifically foods high in saturated fats and free sugars,
is a major cause of the obesity epidemic (Nasser, 2001; Mela, 2001;
WHO, 2013, 2015). Sensory perception and preferences for these
nutrients may contribute to such overconsumption and, hence, it
is important to understand and quantify the extent of interindividual differences.
Fat perception is multi-sensorial, comprising texture, aroma
and taste (Schiffman, Graham, Sattely-Miller, & Warwick, 1998;
Le Calv et al., 2015). The textural aspects of fat are perceived as
Corresponding author at: Department of Food and Nutritional Sciences, The
University of Reading, PO Box 226, Whiteknights, Reading RG6 6AP, UK.
E-mail address: l.methven@reading.ac.uk (L. Methven).
http://dx.doi.org/10.1016/j.foodqual.2016.08.010
0950-3293/ 2016 Elsevier Ltd. All rights reserved.
mouthfeel sensations by trigeminal nerves wrapped around papillae (Whitehead, Beeman, & Kinsella, 1985; Whitehead & Kachele,
1994; Essick, Chopra, Guest, & McGlone, 2003). Greater fungiform
papillae density (FPD) can increase fat texture sensitivity and those
with lower FPD may be less sensitive to the texture of fat (Essick
et al., 2003; Nachtsheim & Schlich, 2013). Hayes and Duffy
(2007) also suggested that subjects with a higher FP count gave
higher creaminess rating to milk-cream mixtures. Additionally,
Hayes, Sullivan, and Duffy (2010) suggested that subjects with
higher FPD had lower preference for high-fat, salty foods.
Nachtsheim and Schlich (2014) demonstrated that subjects with
lower fungiform papillae counts had higher consumption of high
fat milk and spreads than high fungiform papillae counts subjects.
Recently, results from Running, Craig, and Mattes (2015)
demonstrated that medium- and long chain non-esterified fatty
acid (NEFA) were perceived as a unique taste sensation, separate
80
from the other five basic tastes (sweet, bitter, sour, salty and
umami).
PROP taster status is widely used as a marker of taste acuity.
Individuals are commonly classified into three different PROP
taster status groups: supertasters, medium-tasters and nontasters,
according to their relative sensitivity to PROP and sodium chloride
(Tepper, Christensen, & Cao, 2001). A number of studies suggest
that PROP tasters (supertaster and medium-tasters) are not only
more sensitive to PROP solutions, but also perceive higher bitter
intensity from various foods, such as wine (Pickering, Simunkova,
& DiBattista, 2004), caffeinated coffee (Drewnowski, Henderson,
Levine, & Hann, 1999) and grapefruit juice (Drewnowski,
Henderson, & Shore, 1997), as well as to differences in the perception of sweeteners in soft drinks (Zhao & Tepper, 2007). In addition,
authors such as Tepper (2008) have reviewed the role of PROP sensitivity in food choice and nutritional intake. Studies have also suggested that PROP sensitivity is positively associated with fat
perception (Tepper & Nurse 1997, 1998; Hayes & Duffy, 2007;
Nachtsheim & Schlich, 2013). Hayes and Duffy (2007) found subjects with higher intensity ratings towards the bitter-tasting substance PROP gave higher creaminess ratings to high fat cream. It
also has been reported that preference for food with varied fat contents is influenced by PROP taster status. Anliker, Bartoshuk, Ferris,
and Hooks (1991) found that PROP tasters preferred whole milk
significantly more than non-tasters, however, tasters had a significantly lower preference for cheese. Tepper and Nurse (1997)
demonstrated that oiliness and fat ratings were both significantly
higher in PROP taster groups compared to nontasters for salad
dressings of either 10% or 40% fat. Moreover, they found that nontasters preferred the higher fat sample. However, Yackinous and
Guinard (2001) found that PROP taster status was not related to
fat perception in a range of foods (crisps, chocolate drink, milk
pudding and mashed potatoes).
More recent studies have revealed that fat can be detected as a
taste when both mouthfeel and aroma are masked (Mattes, 2009).
Animal (Gilbertson, Liu, Kim, Burks, & Hansen, 2005) and human
(Mattes, 2009; Chale-Rush, Burgess, & Mattes, 2007; Newman &
Keast, 2013; Haryono, Sprajcer, & Keast, 2014; Keast & Costanzo,
2015; Running et al., 2015) studies have suggested that fat might
be the sixth basic taste sensation and some researchers have found
that genotype can influence perception of fat taste. CD36 is responsible for transporting fatty acids and the presence of different single nucleotide polymorphisms (SNPs) of CD36 may influence fat
perception (Laugerette et al., 2005; Abumrad, 2005). Keller
(2012) found that individuals with CD36 A/A genotype (at
rs1761667) rated salad dressing as creamier than either those with
A/G or G/G genotype. Recently, Mrizak et al. (2015) also reported
that obese Tunisian women with CD36 A/A genotype had significantly higher oleic acid detection thresholds. Regarding CD36
genotype and fat preference, Keller (2012) found that the A/A subjects showed higher preference for added fats, oils and spreads,
using data from participants self-report food questionnaires.
Gustin (Carbonic anhydrase VI (CA6)) has been reported to
affect the maintenance and development of taste buds (Henkin,
Martin, & Agarwal, 1999). More recently, studies found that the
polymorphism rs2274333 strongly associated with fungiform
papillae density, where individuals with the G/G genotype had
lower FPD (Melis et al., 2013; Barbarossa et al., 2015). Moreover,
Melis et al. (2013) found that individuals with A/A genotype were
more likely to be PROP supertasters, whereas those with G/G were
more frequently PROP nontasters. As discussed before, fat perception has been reported to associate with papillae density and PROP
taster status. Moreover, considering the role of gustin in fungiform
papillae development and its close association with PROP sensitivity (Padiglia et al., 2010; Calo et al., 2011; Melis et al., 2013), gustin
may plausibly play a role in fat perception.
81
Magnitude Scale (LMS), participants were classified within supertaster, medium-taster and nontaster groups. Data was collected
using Compusense 5 software (Ontario, Canada), for further details
see Shen et al. (2016).
In addition the salt intensity data collected from the
suprathreshold method was used to group participants according
to their rated perception of salt. For each individual their total
score for the three salt solutions was calculated and quartile analysis was carried out on these total scores. Individuals were categorized as low salt perceivers if their total rating was below the value
of the first quartile, medium perceivers if their rating was between
the first and third quartile, and high perceivers if their rating was
equal to or above the third quartile.
2.3.2. Fungiform papillae density
Tongue anterior was coloured using food-grade blue dye (Dr.
Oetker, UK) and two digital photographs were taken from each
participant using a digital microscope (400x magnification, Maplin
Gadget Electronics, UK). A 1 cm2 frame was attached to the end of
the microscope in order that the number of papillae in this area
(number/cm2) could be quantified by two researchers independently. As described by Shen et al. (2016), participants were classified into three groups (low-, medium- and high-FPD).
2.4. Genotypical measurements
Buccal cell samples were collected from participants using sterile Omni swabs (Whatman, UK) and stored in tubes at 20 C prior
to analysis. DNA was extracted using Omega Bio-Tek E.Z.N.A.
Forensic DNA Kit (Norcross, GA, USA) according to the manufacturers instructions. A NanoDrop ND1000 spectrophotometer (Nanodrop Technologies Inc., Rockland, USA) was used to assess DNA
quantity. Concentration of DNA was approximately 15 ng per well.
CD36 (rs1761667) and CA6 (rs2274333) polymorphisms were analysed using the Fluorogenic 50 nuclease assay (TaqMan SNP Genotyping Assay from Applied Biosystems, Foster City, USA).
Genotypes were determined in a 25 lL reaction mix that contained
5 lL DNA sample and 20 lL master mix solution, (6.25 lL DNAse/
RNAse free water, 0.625 lL Taq-Man SNP Genotyping Assay, and
12.5 lL TaqMan Genotyping Master mix (Applied Biosystems Inc.).
The parameters of PCR were as following: 0 min at 55 C followed
by 95 C for 10 min, 92 C for 15 s and 60 C for 1 min (GeneAmp
PCR system 2700, Applied Biosystems). The fluorescence profile
of each well was measured in an Applied Biosystems 7300 System
(Applied Biosystems Inc., Foster City, CA, USA).
2.5. Ice cream formulation and assessment
Four different fat content (6%, 10%, 15% and 20% fat content of
unaerated mix) ice cream samples were produced using the formulations shown in Table 1. All ingredients were homogenised and
pasteurised to 80 C (Boil/5 ice cream pasteuriser, Carpigiani,
Bologna, Italy) before addition of the vanilla flavour and were
refrigerated overnight at 2 C. The mix was then whipped and frozen in a horizontal batch freezer (T. Giusti Ltd, Burton on Trent, UK)
to an overrun of approximately 100% (50 ml air/100 ml finished
product) before depositing into 4 oz (114 ml) paper cups (Bunzl
Catering Supplies, Epsom, UK) and stored at 18 C.
To assess the sensory profile of the ice cream a trained sensory
panel (n = 11) with between 6 months and 4 years experience carried out a quantitative descriptive analysis. A consensus vocabulary of 39 attributes was first reached before scoring each sample
individually, in isolated sensory booths, in duplicate.
To assess the liking of ice cream samples with the study participants, a 9-point hedonic category scale was used for overall liking, liking of taste and liking of mouthfeel. In both the
profiling and liking sessions, sample pots were labelled with 3digit random codes and stored in a 18 C freezer during the sensory session, before serving monadically in a balanced presentation
order. Participants were asked to eat plain crackers and drink
water to remove the creamy aftereffect during one-minute break
between samples.
2.6. Anthropometrics and diet assessment
Body mass index was measured by Tanita body composition
analyser (Tanita BC-418 MA body composition analyser, USA). To
ensure consistency of measurement, all anthropometric measurements, including weight, height, waist and hip circumference, were
carried out by the same researcher. Height was measured in metres
(m) using a wall-mounted stadiometer. Body fat percentage and
weight were measured by Tanita (BC-418 MA body composition
analyser, USA).
In order to assess participants diet, all participants were asked
to complete 133-item Food Frequency Questionnaire (FFQ) (Oxford
food-frequency questionnaire used in European Prospective Investigation of Cancer (EPIC, UK)) based on their last years food consumption. In addition, they completed a 3-day food diary which
was analysed by Diet plan software (Dietplan 6, window version,
Forestfield software, UK) in order to obtain a measure of individual
dietary intake. In order to categorise eating behaviour the 51-item
Three Factor Eating Questionnaire (TFEQ) (Stunkard & Messick,
1985) was used. For restraint, scores of 010 were classified as
low restraint, 1113 high restraint, and 1421 clinical range of
restraint. For disinhibition, scores of 08 were classed as low disinhibition, 911 high disinhibition, and 1216 clinical range of disinhibition. For hunger, scores of 07 were classed as low
susceptibility to hunger, 810 high susceptibility to hunger, and
1114 clinical range of susceptibility to hunger (Harden et al.,
2009).
2.7. Data analysis
All statistical analyses of the study participant data were performed using XLSTAT (version 2015.6.01, France). Data from the
suprathreshold tests was collected on the LMS scale which has
Table 1
The ingredients used to make four different fat levels of ice cream.
Ingredients (g)
10%
15%
20%
625
430
200
37.5
600.1
3107.4
7.5
1041.5
430
200
37.5
570.3
2720.7
7.5
1565
430
200
37.5
532.9
2234.6
7.5
2083.5
430
200
37.5
495.9
1753.1
7.5
Number (N) in Bold was the number of participants in each category. Percentage in Bold was the proportion in the total population (N = 136). n represents the number within each category, and % represents the proportion. BMI
value = Mean SD (standard deviation).
4.2
3.9
3.8
22.9
22.6
23.5
37
37
26
3
3
2
33
59
8
4
7
1
22
52
26
26
60
30
25
50
25
3
6
3
31
63
6
5
10
1
23
50
27
23
51
28
33
50
17
2
3
1
17
43
40
8
21
19
28
56
16
25
49
14
20
44
36
8
18
15
26
55
19
25
52
18
24
52
24
High-FPD
Medium-FPD
Low-FPD
FPD groups
33
70
33
3.5
3.3
5.3
23.5
22.3
23.5
38
25
38
3
2
3
42
0
8
5
0
7
22
28
50
26
32
58
25
25
50
3
3
6
25
69
6
4
11
1
24
51
26
24
52
26
50
33
17
3
2
1
15
52
33
7
25
16
31
49
21
27
43
18
27
54
20
11
22
8
24
48
27
23
46
26
25
50
25
Low perceiver
Medium perceiver
High perceiver
Salt intensity group
34
68
34
5.0
2.8
4.5
23.4
22.3
23.7
26
37
37
2
3
3
33
59
8
4
7
1
28
49
23
32
57
27
25
50
25
3
6
3
38
56
6
6
9
1
28
47
25
28
48
26
17
66
17
1
4
1
31
42
27
15
20
13
26
53
21
23
47
18
15
56
29
6
23
12
34
46
20
32
44
19
28
49
23
Supertaster
Medium-taster
Nontaster
PROP Taster Status
38
67
31
3.6
4.2
3.9
22.4
23.2
23.2
25
50
25
2
4
2
33
59
8
4
7
1
41
54
5
47
63
6
25
67
8
3
8
1
13
81
6
2
13
1
41
52
7
42
53
7
100
0
0
6
0
0
25
62
13
12
30
6
47
50
3
41
44
3
37
51
12
15
21
5
40
56
4
38
53
4
39
54
7
A/A
A/G
G/G
CA6 genotypes
53
74
9
22.9
23.0
22.7
%
38
50
12
3
4
1
n
%
33
33
33
4
4
4
n
%
22
50
28
26
58
32
n
%
8
67
25
1
8
3
n
%
13
31
56
2
5
9
n
%
27
50
23
28
51
23
n
%
33
33
33
2
2
2
n
%
29
42
29
14
20
14
n
%
22
52
26
19
46
23
n
%
17
46
37
7
19
15
27
49
23
%
n
26
47
22
24
49
27
8 (6%)
12 (9%)
116 (85%)
12 (9%)
16 (12%)
102 (75%)
6 (4%)
48 (35%)
88 (65%)
41 (30%)
AGE-2
(3055)
African
Caucasian
East
Asian
West
Asian
Group 1
(12)
Group 2
(34)
Group 3
(58)
(Mean SD)
BMI (kg/m2)
Socio-economic groups
Ethnicity
Male
95 (70%)
Total
(n = 136)
N
%
33
66
37
A/A
A/G
G/G
CD36 genotypes
AGE-1
(1829)
Female
Age
3. Results
Gender
3.0
3.9
4.8
Table 2
Demographic, taste phenotype and taste genotype profiles of participants in the study.
82
83
heterozygous A/G and the remaining 27% were G/G. For gustin
(rs2274333), 39% of the study population had the putatively functional homozygous A/A genotype, more than half of participants
(54%) were heterozygous A/G and the remaining 7% participants
carried the G/G genotype.
According to the intensity rating from PROP suprathreshold
test, 28% of participants were supertasters, 49% were classified as
PROP medium-tasters and the remaining 23% participants were
nontasters. From quartile analysis of the salt intensity data, participants were grouped into perception groups according to their
overall ratings the three salt solutions: high salt perceivers (HSP,
total rating 129233), medium (MSP, total rating 58128) and
low (LSP, total rating 1857). Participants were also classified into
three FPD groups. Twenty-four percent participants were in the
high-FPD group (FPD range from 76 to 132 papillae/cm2), over half
of participants (52%) were in the medium-FPD group (FPD range
from 43 to 73 papillae/cm2) and 24% participants were the lowFPD group (FPD range from 12 to 42 papillae/cm2).
Table 3
Categorization of subjects from Three Factor Eating Questionnaire (TFEQ).
Categories
Score range
Mean SD
Low restraint
High restraint
Clinical range of restraint
87
21
28
64
15
21
010
1113
1421
63
12 1
16 2
Low disinhibition
High disinhibition
Clinical range of disinhibition
110
16
10
81
12
7
08
911
1215
52
10 1
13 1
109
21
6
80
15
5
07
810
1111
42
91
11 0
n presented the number of participants in each category, and % represents the proportion in the total population (N = 136).
5
Overall Liking
6%
10%
Liking of taste
15%
Liking of Mouthfeel
Fig. 1. Participant mean liking of four ice cream samples varying in fat content. Error bars represent standard error of the mean (N = 136).
84
mouthfeel substantially lower for the 20% fat ice cream compared
to their rating for other three samples (shown in Supplementary
Tables S1S3). Similarly, no effect of salt intensity group was found
in overall liking (p = 0.19), liking of taste (p = 0.32) and liking of
mouthfeel (p = 0.81). However, there was a significant interaction
between ice cream samples and salt intensity group for overall liking (p = 0.03), liking of taste (p = 0.02) and liking of mouthfeel
(p = 0.03). The LSP group rated their overall liking, liking of taste
and mouthfeel substantially higher for the 20% fat ice cream compared to the MSP (p = 0.017, p = 0.047 and p = 0.55 respectively)
and HSP (p = 0.12, p = 0.068 and p = 0.32 respectively) groups
(Fig. 3) (detail in Supplementary Tables S1S3).
For FPD group, no effect was found in overall liking (p = 0.063),
liking of taste (p = 0.11) and liking of mouthfeel (p = 0.11). There
were no significant interactions between ice cream samples and
FPD group, however there was a trend for the Low-FPD group to
rate their overall liking, liking of taste and mouthfeel lower for
the 20% fat sample (shown in Supplementary Tables S1S3).
Comparing the different CD36 genotypes, there was also no significant effect on overall liking (p = 0.24), liking of taste (p = 0.37)
nor liking of mouthfeel (p = 0.60) of the ice cream. There were no
80.0
70.0
60.0
50.0
40.0
30.0
20.0
10.0
0.0
6%
10%
15%
Fig. 2a. Appearance and mouthfeel attributes found to differ significantly between ice cream samples formulated to different fat contents. Error bars represent +/ half the
Tukey HSD values. AE = aftereffect, evaluated 30 s post swallowing. Appearance attributes not found to differ significantly between samples were: glossy, sticky, density in
pot. Mouthfeel attributes not found to differ significantly between samples were: mouthdrying and mouthdrying aftereffect.
50.0
45.0
40.0
35.0
30.0
25.0
20.0
15.0
10.0
5.0
0.0
Dairy Fat
Odour
6%
Cardboard
Odour
10%
Acid (sour)
Taste
15%
20%
Dairyfat
(creamy)
Flavour
O Flavour
(Plasc or
Chemical)
Bier AE
Fig. 2b. Odour, taste and flavour attributes found to differ significantly between ice cream samples formulated to different fat contents. Error bars represent +/ half the
Tukey HSD values. AE = aftereffect, evaluated 30 s post swallowing. Odour attributes not found to differ significantly between samples were: vanilla, sweet, caramel, milk
powder, sour milk, overall odour intensity and off odour. Taste and flavour attributes not found to differ significantly between samples were: sweet, bitter, vanilla, caramel
and cooked milk, sweet aftereffect and sour aftereffect.
85
(i)
8.0
7.5
7.0
6.5
6.0
5.5
5.0
4.5
4.0
3.5
3.0
6%
10%
15%
20%
(ii)
8.0
7.5
7.0
6.5
6.0
5.5
5.0
4.5
4.0
3.5
3.0
6%
10%
15%
20%
Table 4
Mean of overall liking ratings of participants grouped by AHC.
Cluster
6% Fat
10% Fat
15% Fat
20% Fat
Cluster 1: nondiscriminator
Cluster 2: high-fat disliker
Cluster 3: high-fat liker
71
41
24
52%
30%
18%
7.6 0.1aA
6.5 0.2bcBC
4.5 0.4dD
7.7 0.1aA
6.1 0.3bcBC
5.6 0.4cdCD
7.5 0.2aA
6.9 0.3abAB
6.8 0.3abcABC
7.4 0.1aA
2.6 0.2eE
6.8 0.3abcABC
n = number of participants in each group, and % present the proportion in total number of participants (N = 136). Numbers in table represent the mean of overall liking rating
of each group for four ice cream samples. acMean values not sharing a common superscript lowercase letter in the same row differed significantly (p < 0.05) in same row. A
Mean values not sharing a common superscript uppercase letter in the same column differ significantly (p < 0.05).
86
90
(i)
85
a
80
75
70
ab
b
65
60
55
50
45
40
Supertaster
Medium-taster
Non-taster
(ii) 40
Fat Intake (% of Total Energy Intake)
Altering the fat content of the ice cream samples did alter the
sensory profile of the ice creams. Of 39 attributes assessed, 20 differed significantly between samples (Figs. 2a and 2b).
The sensory profile results conclude that the ice creams formulated to different fat contents differed in their sensory profile. The most striking differences in mouthfeel were, as
expected, that increasing fat content increased body (p = 0.01),
smoothness (p < 0.0001), mouthcoating (p = 0.01), creaminess
of mouthfeel (p < 0.0001), was slower to dissolve in mouth
(p = 0.0002) and led to greasiness on the lips (p = 0.04). The
6% fat formulation led significantly to more ice crystals being
felt in the mouth than compared to the 10% and 20% fat samples (sample effect p = 0.0005) and was perceived to be more
powdery than the 15% and 20% fat samples (p = 0.0001).
Increasing the cream in the ice cream formulation did lead to
an increase in dairy fat (creamy) flavour (p = 0.002) where the
6% and 10% fat ice creams were significantly lower than the
15% and 20% fat samples. The lower fat samples tended to be
slightly higher in off notes; the 6% fat sample being significantly higher in off-flavour than the 15% and 20% fat samples.
Fat appeared to mask acidic taste and bitter aftertaste in the
ice cream as these were significantly higher, although still
scored at low intensity, in the 6% fat sample compared to the
15% and 20% fat samples.
35
30
25
20
Liking Cluster 3 (high fatlikers, n=24)
Fig. 4. Mean fat intake (g) per day calculated from 3 day diet diaries; (i) differences
between PROP taster status groups (g/day), (ii) differences between ice cream liking
cluster groups (fat as % total energy intake/day). Error bars represent standard error
of the mean for each group. abBars with different letters indicate significant
difference at p < 0.05.
energy from fat (%) (p = 0.031), where the high restraint group
tended to have a substantially lower percentage fat intake than
the low restraint group (28% versus 33%, p = 0.056).
A significant difference in BMI was found between disinhibition
groups (p = 0.001), where the clinical range of disinhibition group
had significantly higher BMI than low disinhibition group (26.6
versus 22.5 kg/m2, p = 0.002)). There was no significant difference
in BMI between restraint groups (p = 0.51) nor susceptibility to
hunger groups (p = 0.18).
500
450
400
350
300
250
200
150
100
50
0
Liking Cluster 1 (Non- Liking Cluster 2 (high Liking Cluster 3 (high
discriminators, n=71) fat-dislikers, n=41)
fat-likers, n=24)
Fig. 5. Mean intake of dairy products (g per day) calculated from food frequency
questionnaire. Error bars represent standard error of the mean for each group.
ab
Bars with different letters indicate significant difference at p < 0.05.
87
4. Discussion
In this study, both genotypes (CD36 (rs1761667) and CA6
(rs2274333)) and phenotypes (PROP taster status, salt sensitivity
groups and FPD group) of taste sensitivity measurements were
used to classify participants. Although PROP taster status is the
most common method to classify participants according to taste
sensitivity, it is a reflection on both overall taste sensitivity and
specific sensitivity to the thiourea group in PROP (Hayes & Keast,
2011), which varies with genotype for the bitter taste receptor
T2R38. As taste sensitivity to fat will not be effected by the specific
sensitivity to the thiourea group, we decided to separately take the
salt intensity measures from the PROP taster status test as a marker of general taste sensitivity (as described in Section 2.3). Considering the relationship between the genotype and phenotype of
taste sensitivity, the results concluded that there is no significant
association between CD36 and CA6. Moreover, no significant association was found between CD36 and either PROP taster status or
salt intensity group. Additionally, there was a lack of relationship
between CD36 and FPD groups in the current study. Gene expression results from animal (Laugerette et al., 2005; Fukuwatari
et al., 1997) and human (Simons, Kummer, Luiken, & Boon, 2011)
studies have found CD36 to be more associated with circumvallate
papillae than fungiform papillae, so the lack of relationship with
FPD was perhaps not surprising. Our results did not find any relationship between CA6 (gustin) genotype and FPD groups, nor
between CA6 and PROP taster status or salt taste intensity group.
Similar results were also found in Feeney and Hayes (2014) study,
as discussed in Shen et al. (2016).
The results from our study conclude that the taste genotypes
(CD36 and CA6) and phenotypes (PROP taster status and FPD
group) did not significantly affect liking ratings for ice cream samples varying in fat content. However, there was a trend for both
PROP non-tasters and the low-FPD group to show substantially
lower liking of the 20% fat content sample and, conversely, there
was a significant interaction between ice cream samples and salt
intensity group where the low salt perception (LSP) group rated
their liking (overall, taste and mouthfeel) substantially higher for
the 20% fat ice cream. From results of previous studies we predicted those of low taste sensitivity would like the higher fat sample more. A previous study by Tepper and Nurse (1997) found that
PROP supertasters (n = 25) and medium-tasters (n = 25) were able
to discriminate different in fat content 40% and 10% salad dressing,
but nontasters (n = 25) could not. Moreover, they also suggested
that nontasters showed higher hedonic preference for the 40% fat
salad dressing despite not being able to discriminate the difference
in fat content between the two samples (Tepper & Nurse, 1998).
Additionally, Keller, Steinmann, Nurse, and Tepper (2002) also
found PROP nontaster children preferred full-fat milk compared
to PROP tasters. Although this is in-line with the results from our
salt groupings, it contradicts those from our PROP and FPD groupings. However the sensory profiling results demonstrated that the
change in fat content led to numerous differences between the ice
cream samples. Although the higher fat samples were more
smooth, coating and creamy in mouthfeel, as well as higher in
dairy fat flavour, they were also less powdery and lower in ice crystals in the mouth. Therefore, the PROP supertaster and high FPD
group results, where mean liking was not significantly different
between the low and high fat content samples, are perhaps less
surprising. Higher sensitivity to the ice and powdery mouthfeel
in the low fat samples might have reduced liking for some of these
participants; but also their higher sensitivity to the coating and
creamy mouthfeel in the high fat samples might reduce liking for
other supertasters. Hence leading to no overall effect on the mean
liking results for these groups. Overall therefore, although difference in mouthfeel and taste perception have influenced the liking
88
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