Food Quality and Preference 31 (2014) 19

Contents lists available at ScienceDirect

Food Quality and Preference

journal homepage: www.elsevier.com/locate/foodqual

Aromataste interactions between a model cheese aroma and ve basic

tastes in solution
Jun Niimi a,b, Andrew I. Eddy b, Amy R. Overington c, Samuel P. Heenan a, Patrick Silcock a, Phil J. Bremer a,,
Conor M. Delahunty d
a

Sensory Science Research Centre, Department of Food Science, University of Otago, Dunedin, New Zealand
CSIRO Animal, Food and Health Sciences, North Ryde, NSW, Australia
Fonterra Research and Development Centre, Palmerston North, New Zealand
d
CSIRO Animal, Food and Health Sciences, Adelaide, SA, Australia
b
c

a r t i c l e

i n f o

Article history:
Received 21 January 2013
Received in revised form 21 May 2013
Accepted 26 May 2013
Available online 12 July 2013
Keywords:
Cross-modal sensory interactions
Cheese avour intensity
Taste intensity
Enhancement
Suppression

a b s t r a c t
The avour perception of cheese results from complex sensory interactions between tastes and aromas.
Using a model cheese solution, this study investigated perceived interactions between each of ve basic
tastes and a cheese aroma mixture containing ten volatile compounds commonly found in cheese. The
ve tastes sucrose (sweetness), sodium chloride (NaCl) (saltiness), monosodium glutamate (MSG)
(umami), lactic acid (sourness), and caffeine (bitterness) were individually mixed with cheese aroma
in water using a 5 taste level (0.2 log series) by 3 aroma level (0.5 log series) design. Aroma controls with
no added taste were also included. This resulted in 18 samples for each single tastearoma combination.
An additional 18 samples were produced using a mixture of all 5 tastes with the 3 aroma levels. A panel of
trained assessors (n = 10) evaluated cheese avour intensity and taste intensity using 100 point line scales.
Evaluation was carried out in duplicate, with samples grouped by taste type; 1 evaluation session per
taste per replicate. Within type, order of presentation was balanced, and taste type order was randomised
between replicates. Cheese avour intensity was enhanced by sucrose and NaCl, while being suppressed
by lactic acid. NaCl enhanced cheese avour intensity the most at high aroma level, while lactic acid suppressed the most at low aroma level. When MSG level was increased, cheese avour intensity was
enhanced at both low and medium aroma levels, but was suppressed at the high aroma level. The greatest
enhancement of cheese avour intensity was found with the mixture of 5 tastes. Aroma signicantly
enhanced umami and bitterness, but did not enhance sweetness, saltiness, or sourness. This study
showed that the perceived interaction between taste and cheese aroma depended on taste type and on
the concentration levels of both taste type and aroma. The mixture of tastes was more effective at
enhancing cheese avour intensity than single tastes. This study provides knowledge that will underpin
further study of tastearoma interactions in a model cheese that aims to optimise cheese avour intensity and character.
2013 Elsevier Ltd. All rights reserved.

1. Introduction
Flavour is of key importance to consumers acceptance of
cheese. The avour perception of cheese is inuenced by multiple
sensory modalities, before and during consumption, including
appearance, taste, aroma, chemical irritation, and texture (Delahunty & Drake, 2004).
Aroma perception occurs by olfaction: aroma active compounds
stimulate olfactory receptors situated on the roof of the nasal
cavity. In the context of avour perception during consumption,

Corresponding author. Tel.: +64 3 479 5469.

E-mail address: phil.bremer@otago.ac.nz (P.J. Bremer).
0950-3293/$ - see front matter 2013 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.foodqual.2013.05.017

aroma compounds are perceived retronasally (Rozin, 1982). Many

different types of aroma compounds are responsible for the aroma
of cheeses, including short to medium chain fatty acids, aldehydes,
alcohols, ketones, esters, and sulphur compounds (Curioni & Bosset, 2002). The aroma character of a cheese is determined by the
combination of aroma compounds and is not characteristic of
any single compound (Zehentbauer & Reineccius, 2002). Different
cheeses can share many of the same aroma compounds, but their
varying proportion differentiates one type of cheese from another
(Engels, Dekker, de Jong, Neeter, & Visser, 1997).
Taste perception occurs during gustation as taste active compounds stimulate taste receptors on the tongue. Taste perception
can be separated into ve basic taste characters, including sweetness, saltiness, umami, sourness, and bitterness, all of which are

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

present in the taste of cheeses. Compounds identied as contributing to the basic taste characters of a variety of cheese types include
NaCl for saltiness, organic acids for sourness, amino acids (particularly glutamate) for umami, hydrolysed peptides with hydrophobic
amino acids and mineral salts contributing to bitterness, and salt
propionates and amino acids contributing to sweetness (Andersen,
Ard, & Bredie, 2010; Engel, Septier, Leconte, Salles, & Le Qur,
2001; Kubckov & Grosch, 1998; Salles, Septier, Roudotalgaron,
Guillot, & Etievant, 1995; Salles et al., 2002; Taborda et al., 2008;
Toelstede & Hofmann, 2008a, 2008b; Warmke, Belitz, & Grosch,
1996). However, the relative contribution of each taste character
towards the overall cheese taste perception is not well known.
The combined stimulation of aroma and taste inuences the
perceived intensity of each other, despite gustatory and olfactory
receptors and their neural pathways to the brain being physiologically separate. The resultant effect is a cross-modal sensory interaction, and can result in an enhancement or suppression of
attribute intensities (Noble, 1996). For enhancements to occur, it
is believed that the taste and aroma must be harmonious or congruent with each other (Schifferstein & Verlegh, 1996). It is possible for congruency to arise from learning through repeated
exposure to a particular aroma and taste combination (Prescott,
2012). During the perception of a congruent mixture of taste and
aroma, it is thought that cross modal interaction is integrated in
specic areas of the brain. The orbitofrontal cortex, the anterior
cingulated cortex, and the insula have been shown to be the important areas for integration (Eldeghaidy et al., 2011; Small & Prescott,
2005; Small et al., 2004). Congruent mixtures of aroma and taste
show consistent enhancements across replicate measurements
(Green, Nachtigal, Hammond, & Lim, 2012). Mixtures of stimuli
that are incongruent can either show no change in perception from
when the stimuli are individually perceived, or suppression (Prescott, 1999; Stevenson, Prescott, & Boakes, 1999).
A number of studies have reported that perceived taste, including sweetness, bitterness, saltiness, and sourness, can be enhanced
by aroma (Bonnans & Noble, 1993; Caporale, Policastro, & Monteleone, 2004; Djordjevic, Zatorre, & Jones-Gotman, 2004). In the context of cheese, cross-modal interactions have not received much
research attention. Recently, with salt reduction as the aim, it
was demonstrated that perceived saltiness of both a solution,
and a model cheese matrix representing mozzarella, could be enhanced by the addition of specic aroma characters, including
Comt cheese, goats cheese, Roquefort cheese, or a combination
of pure compounds commonly found in cheese (Lawrence, Salles,
Septier, Busch, & Thomas-Danguin, 2009; Lawrence et al., 2011;
Pionnier et al., 2004). There is limited knowledge on how taste
characters such as sweetness, sourness, umami, and bitterness
inuence perceived cheese avour. Early work revealed that the
combination of tastes as a whole contributes signicantly to the
avour intensity of cheeses (Aston & Creamer, 1986; McGugan,
Emmons, & Armond, 1979). No further reports have been shown
that follow through with these ndings.
The objective of the current study was to investigate crossmodal sensory interactions between each of the ve basic taste
types and a cheese aroma. All 5 basic taste characters, sweet, salty,
sour, umami, and bitter, were tested with a complex cheese aroma
consisting of 10 compounds. Each character was tested individually, and then all 5 tastes were mixed with aroma.

sucrose (Bundaberg Sugar, Spring Hill, QLD, Australia), sodium

chloride (NaCl) (Sigma Aldrich, Sydney, NSW, Australia), monosodium glutamate (MSG), DL-lactic acid (85%, FCC Kosher, food grade
SAFC supply solutions, Australia), and caffeine (SigmaAldrich),
respectively. The MSG was prepared by mixing sodium hydroxide
(98%) (Sigma Ultra, food grade: SigmaAldrich) with L-glutamic
acid (98.5%, FCC Kosher, food grade: SAFC, Sydney, NSW, Australia)
in water. Aroma compounds were 2-butanone, 2-heptanone,
2-nonanone, diacetyl, ethyl butyrate, butyric acid, methional,
3-methylbutanal (all obtained from Givaudan Australia Pty Ltd,
Sydney, NSW, Australia), ethyl hexanoate (Firmenich Ltd, Balgowlah, NSW, Australia), and 3-methylbutanoic acid (SigmaAldrich).

2.2. Experimental design

The ve tastes; sucrose, NaCl, MSG, lactic acid, and caffeine,
were individually mixed with cheese aroma in water using a 5 level of taste by 3 level of aroma design (Tables 1 and 2). For each
taste and aroma combination, taste controls at each of the three
aroma levels were also prepared; these samples were aroma
solutions that did not contain taste, meaning that the same taste
control samples were always used. This resulted in 18 samples
for each single-taste and aroma combination, also using the same
design as above. The taste concentration levels were chosen to represent the taste intensity range for each type, typically found in
foods. The highest concentrations for each type were matched by
panel intensity ratings. The concentration ranges for each of these
tastes were obtained by reducing the concentrations in 0.2 log
steps four times from the highest concentration. A solution containing a mixture of all ve tastes was also combined with the aroma. The tastes were mixed at equal concentration levels to when
compared singly, as shown in Table 1. For example at taste level 1,
the levels of all ve tastes in that row of Table 1 were combined
together. This gave a mixture of sucrose, NaCl, MSG, lactic acid,
and caffeine at concentrations of 1.9%, 0.13%, 0.07%, 0.07%, and
0.03%, w/v, respectively. This was repeated at other taste levels
to give levels 2, 3, 4, and 5.

Table 1
Five concentrations of ve taste types used for the aromataste interactions; controls
of each taste type were 0% (w/v).
Sucrose (%,
w/v)

NaCl (%,
w/v)

MSG (%,
w/v)

Lactic acid
(%, w/v)

Caffeine (%,
w/v)

1
2
3
4
5

1.90
3.01
4.78
7.57
12.00

0.13
0.20
0.32
0.50
0.80

0.07
0.11
0.18
0.28
0.45

0.07
0.11
0.18
0.28
0.45

0.03
0.05
0.08
0.13
0.20

Table 2
Concentrations of 10 aroma compounds that represent cheese aroma.

2. Materials and methods

2.1. Samples
The taste types used to represent pure taste characters of
sweetness, saltiness, umami, sourness, and bitterness and were

Tastant
level

Compounda

CAS-No.

2-Butanone
2-Heptanone
2-Nonanone
3-Methylbutanal
3-Methylbutanoic acid
Butyric acid
Diacetyl
Ethyl butyrate
Ethyl hexanoate
Methional

78-93-3
110-43-0
821-55-6
590-86-3
503-74-2
107-92-6
431-03-8
105-54-4
123-66-0
3268-49-3

Aroma level (lg/L)

Low

Medium

High

0.200
0.120
0.040
0.020
0.060
0.800
0.200
0.120
0.040
0.020

0.630
0.380
0.130
0.060
0.190
2.530
0.630
0.380
0.130
0.060

2.000
1.200
0.400
0.200
0.600
8.000
2.000
1.200
0.400
0.200

Levels increased by 0.5 log series, in an equal proportion for all compounds.

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

The aroma compounds were chosen to provide a cheese-like

character, with their relative concentration levels being based on
headspace analysis of a model cheese produced by Overington
et al. (2010) (Table 2). Their mixtures resembled a Cheddar cheese
like aroma. The concentration range for aroma was achieved by
obtaining the maximum concentration and reducing the concentrations of all of the aroma compounds in 0.5 log steps twice to
give three aroma levels.
All samples were evaluated in duplicate, with samples grouped
by taste type; one evaluation session per taste per replicate. Within
each taste type, the order of sample presentation was balanced
using CycDesigN Ver. 2.0 (CSIRO Forestry and Forest Products, Canberra, ACT, Australia) to ensure that each sample had an equal
chance of being presented at any position to the assessors. Samples
were blind coded with three digits. Taste type order was also randomised between replicates.
2.3. Sensory procedure
A panel of experienced assessors (n = 10) evaluated the samples.
These assessors had been screened in accordance with ISO standards and had previously been trained in the use of scales with a
wide range of food products and basic taste discrimination. While
assessors had extensive prior experience with sensory evaluation,
they did not undergo training specic to this study. Rather, assessors were exposed to aroma and taste separately prior to the rst
session, to inform them of the perceptual characters. Exposure to
single taste solutions including sweetness, saltiness, umami, sourness, and bitterness was repeated at the beginning of every session,
with the highest concentration of taste being presented. The highest concentration of each taste was assigned an intensity of 80 on a
100 point intensity scale. The intensity of 80 points was chosen so
that aroma enhancement of taste could be measured. For each
taste solution, the assessors were informed of the single taste
character.
On each day of testing, assessors were presented with samples
in three sets; each set consisted of six samples. To prevent sensory
fatigue, breaks of 15 min were scheduled between sample sets. The
samples were presented at ambient temperature in booths under
white light and positive air pressure. The assessors were instructed
to taste and swallow the sample and then rate the intensity of
cheese avour, followed by the intensity of the taste attribute(s)
specic to the particular session. The intensity of cheese avour
in this case was a measure of the holistic perception of cheese avour, including taste and aroma. Ratings were made on an unstructured 100 point scale, which was anchored at 5 and 95 points with
low and high, respectively. When evaluating the samples with
a mixture of ve tastes, assessors rated the intensities of six attributes; cheese avour intensity, sweetness, saltiness, umami, sourness, and bitterness. Unsalted water crackers, cucumber, and water
were used as palate cleansers between each sample. Data were acquired using Compusense ve, Ver. 4.6 (Compusense Inc., Guelph,
ON, Canada).
2.4. Data analysis
Differences in cheese avour intensity compared with taste
controls (aroma only in water) were calculated for all samples
for each assessor. Differences were calculated separately at each
aroma level, resulting in 15 values per taste type per replicate.
By analysing the data in this way, it accounted for differences
not only within taste type but also in overall level between tastes
types due to a contrast effect observed, whilst maintaining aroma
level as a variable.
A one-way analysis of variance (ANOVA) was carried out to
compare relative enhancement or suppression of cheese avour

intensity between taste types, including all aroma levels.

Fishers least signicant difference (LSD) post hoc test was
used where signicant differences were found at 5% alpha level.
A Students paired t-test, at 5% alpha level, was carried out to
determine if a taste type caused statistically signicant
enhancement or suppression compared with the control with
aroma levels combined.
A univariate general linear model (GLM) ANOVA was carried out
for each taste type separately. Panel means of difference values of
cheese avour intensity and unadjusted taste intensities were taken as the dependent variables. The aroma level, taste level, and
replicate were taken as the independent variables for analysis of
both types of dependent variables, while assessors were taken as
a random variable. Two-way interactions were also investigated,
including taste level  aroma level, taste level  replicate (for differences in cheese avour intensity only), and taste level  aroma
level, aroma level  replicate (for taste intensity only). Interactions
with replicate were analysed for the stability of cross-modal sensory interactions towards cheese avour and taste intensity ratings. Fishers LSD post hoc test, at 5% alpha level, was used
where signicant differences were found for all attribute intensities. An additional students paired t-test at 5% alpha level was carried out within taste type, with aroma levels combined to
determine whether each level of taste either enhanced or suppressed cheese avour intensity compared with its corresponding
control.
A one-way ANOVA was carried out to compare the aroma controls of each taste type to investigate contrast effects. The mean
cheese avour intensities were taken as the dependent variable
and the taste type as the independent variables.
A students t-test at 5% alpha level was carried out to compare
the intensities of each taste attribute between each single taste
solution and the mixture of ve tastes. This was carried out to
investigate the degree of taste suppression in the mixed taste,
and was performed at each taste concentration.
ANOVA was also carried out to determine the effect of aroma level on the unadjusted cheese avour intensity for each taste type.
In this case panel mean avour intensity scores were analysed. All
statistical analyses were performed using the statistical software
SPSS Ver. 17 (SPSS Inc., Chicago, USA).

3. Results and discussion

3.1. Effect of tastes on cheese avour intensity
The overall difference in cheese avour intensity of samples
containing either a single taste and cheese aroma, or a mixture of
5 tastes and cheese aroma, was assessed against a control sample
containing only cheese aroma (Fig. 1). Compared with the control,
avour intensity was signicantly enhanced by the addition of sucrose or NaCl, and signicantly suppressed by the addition of lactic
acid. The addition of caffeine or MSG did not cause a signicant difference in cheese avour intensity compared with the control. In
addition, cheese avour intensity was signicantly enhanced by
the mixture of ve tastes, and this enhancement was greater than
the enhancement due to the addition of either sucrose or NaCl. The
nding that the addition of a mixed taste to aroma enhanced avour intensity more than the addition of a single taste is in agreement with previous studies that investigated interactions between
mixed tastes and aroma (Nasri, Septier, Beno, Salles, & ThomasDanguin, 2013; Pfeiffer, Hort, Hollowood, & Taylor, 2006). In one
of these previous studies, the strawberry avour intensity of a mixture of sucrose, organic acid, and aroma was more than double the
intensity of mixtures of either taste singly and aroma (Pfeiffer
et al., 2006).

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

Mean difference in cheese flavour intensity

20
***
e

15
***
d

***

10

***
a

Lactic acid

Caf f eine

-5

-10
MSG

Sucrose

NaCl

Mixture

Tastant type

Fig. 1. Mean differences in cheese avour intensity ( standard error), compared

with a taste-free control, across taste types including single tastes and a mixture of
ve basic tastes. Means with the same superscripts are not signicantly different
according to Fishers LSD. Signicant difference from control: = p < 0.001.

The effect of taste concentration on cheese avour intensity was

examined for each taste type and for the mixture of ve tastes
(Fig. 2). The effect of aroma level was included in the analysis. Sucrose concentrations of 3.01% and above signicantly enhanced
cheese avour intensity compared with the control (Fig. 2a), with
an enhancement of up to 9.6 points at 4.78% sucrose. This result

(a)

Difference in cheese flavour

intensity

20

**

15

10

5
0
-5

-10
-15
-20
1.90

3.01

4.78

7.57

showed that there was a minimum concentration below which sucrose did not enhance cheese avour intensity, and a maximum
concentration above which further enhancement did not occur. A
signicant sucrose  aroma interaction was not detected, showing
that the cheese avour enhancement due to increasing sucrose
concentration was the same across all aroma concentrations.
Sweet character is important to the avour of Swiss cheese due
to the presence of salt propionates (Preininger, Warmke, & Grosch,
1996; Warmke et al., 1996). Sweetness has also been reported to
contribute somewhat to the avour of Gouda, due to the presence
of sweet tasting amino acids (Toelstede & Hofmann, 2008a). Sweetness is thought not to be important to Cheddar cheese avour
(Andersen et al., 2010; Toelstede & Hofmann, 2008a).
The results for NaCl were similar to those found for sucrose,
with concentrations of 0.2% NaCl and above signicantly enhancing cheese avour intensity compared with the control (Fig. 2b),
and an enhancement of up to 11.7 points at 0.5% NaCl. No signicant interaction of NaCl concentration  aroma concentration was
detected, showing that the cheese avour enhancement due to
increasing NaCl concentration was the same across all aroma concentrations. The enhancement of cheese avour intensity by NaCl
was expected, as NaCl is known to contribute towards the perceived avour intensity of Cheddar cheese (Schroeder, Bodyfelt,
Wyatt, & McDaniel, 1988). The NaCl most likely did not cause a
salting-out effect on the volatile compounds that may have contributed to the perceived enhancement, as volatile compounds

20
Difference in cheese flavour
intensity

***

15

10
5
0
-5
-10
-15
-20

12.00

0.13

0.20

Sucrose (% w/v)

10
5
0
-5

-10
-15
-20
0.11

0.18

0.28

(d)

10
5

0.28

0.45

-5

-15
-20
0.07

20
Difference in cheese flavour
intensity

Difference in cheese flavour

intensity

**

-10

0.45

(e)

15
10
5
0
-5
-10
-15
-20
0.08

**

0.11

0.18

Lactic acid (% w/v)

20

0.05

0.80

15

MSG (% w/v)

0.03

0.50

20
Difference in cheese flavour
intensity

Difference in cheese flavour

intensity

15

0.07

0.32
NaCl (% w/v)

(c)

20

(b)
**

0.13

Caffeine (% w/v)

0.20

15

***

***

***

3
Tastant level

(f)
*

**

10
5
0
-5
-10
-15
-20
1

Fig. 2. Mean differences in cheese avour intensity ( standard error) from the control. Tastants used were (a) sucrose, (b) NaCl, (c) MSG, (d) lactic acid, (e) caffeine, and (f)
mixture of ve tastes (levels according to Table 1). Means with the same superscripts are not signicantly different according to Fishers LSD. Signicant difference from
control: = p < 0.05, = p < 0.01, and = p < 0.001.

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

Difference in cheese flavour

intensity

20
15
10
5
0
-5

-10
-15
-20
0.07

0.11

0.18

0.28

0.45

MSG (% w/v)

Fig. 3. Mean differences in cheese avour intensities ( standard error), from each
aroma level control, at three aroma levels by MSG (white bars = low aroma, grey
bars = medium aroma, and black bars = high aroma).

have been shown to start salting-out at 5% NaCl with the degree of

salting-out depending on the volatile compound (Yang & Peppard,
1994). With NaCl increasing to a maximum concentration of 0.8%
in the current study, the concentration was unlikely to be high enough to salt volatile compounds out.
Overall, MSG did not enhance cheese avour intensity compared with the control (Fig. 2c). There was a signicant (p < 0.05)
MSG concentration  aroma concentration interaction. Cheese
avour intensity was enhanced by MSG at low aroma level,
unchanged at medium aroma levels, and suppressed at the high aroma level (Fig. 3). A previous study showed avour intensity
enhancement when umami tastes were perceived together with
savoury aroma, along with increased brain activity when measured
with functional magnetic resonance imaging (McCabe & Rolls,
2007). Umami tastes have been reported to have complex interaction effects, which can be dependent on specic attributes. There
has been evidence of enhancement of nutty, cocoa, and potato avour intensities in a model broth (Ventanas, Mustonen, Puolanne,
& Tuorila, 2010), and suppression of beef, green onion, and garlic
avour intensities in beef soup (Jung, Hong, & Kim, 2010). In the
current study, the sensory interaction effects induced by MSG were
dependent on specic aroma levels, conrming the complex nature
of umamiaroma interactions towards avour perception.
Higher concentrations of lactic acid (0.28% and 0.45%) signicantly suppressed cheese avour intensity compared with the control (Fig. 2d); suppression of 9.1 points at 0.28%, and suppression of
10.2 points at 0.45% lactic. These two highest lactic acid levels
signicantly (p < 0.01) suppressed the cheese avour more than
the other lactic acid levels. No signicant interaction of lactic acid
concentration  aroma concentration was detected, showing that
the cheese avour suppression due to increasing lactic acid concentration was the same across all aroma concentrations. Suppression of avour with increasing acidity has been previously reported
in yoghurt. A reduction in pH caused suppression of sweet, buttery,
and creamy avour attributes (Ott, Hugi, Baumgartner, & Chaintreau, 2000). Physico-chemically, it has been previously shown that
a reduction in pH from 4 to 2 using citric acid in soft drinks decreases the headspace concentration of limonene by approximately
20 ng/ml, while very little change in headspace concentration was
reported for esters (Hansson, Andersson, Leufvn, & Pehrson,
2001). However with the total range of acid used in the current
study being much less than Hansson et al. (2001), the impact of
pH inducing a marked change in volatile release to in turn inuence perception would be minimal. The suppression of cheese
avour intensity may have been due to the sourness overpowering
the cheese avour perception.
Overall, caffeine did not enhance cheese avour intensity
compared with the control (Fig. 2e). No signicant interaction of

caffeine concentration  aroma concentration was detected, where

no change in cheese avour intensity was detected with increasing
caffeine concentration across all aroma concentrations.
The mixture of ve tastes signicantly enhanced cheese avour
intensity at all taste levels compared with the control (Fig. 2f). The
greatest enhancement compared with the control was found at
taste levels 2, 3, and 4, where magnitudes of enhancement were
15.5, 14.0, and 14.7, respectively. This nding suggested that there
was an optimum range for avour enhancement. No signicant
interaction of taste concentration  aroma concentration was detected, suggesting that the enhancement of cheese avour intensity with increasing concentrations of mixtures of tastes were the
same across all aroma concentrations. The high contribution of
mixed tastes to cheese avour intensity demonstrated that a mixture of tastes was more effective in inducing an enhancement than
a single taste. In addition, the magnitude of enhancement was
greater at low concentrations of the taste mixture compared with
that of single tastes. However, increasing concentrations of the
mixture of tastes did not further enhance the cheese avour intensity, possibly because the taste intensity became too high, dominating the aroma character, and was no longer cheese-like.
There were no signicant replicate  taste interactions towards
the difference in cheese avour intensity for any of the single taste
types, or for the mixture of ve tastes. This suggested that the
cheese avour intensity measurements for each taste type were
stable between replicate measurements, demonstrating stability
of the cross-modal sensory interaction effects.
The control samples, which contained aroma only, were identical samples across all taste types tested across sessions, yet the
cheese avour intensity was signicantly (p < 0.001) different
when tested amongst different taste types . The aroma only samples tested amongst the ve tastants mixture were perceived as
signicantly lowest in mean cheese avour intensity (20.9). The
aroma only samples tested amongst NaCl (32.5), sucrose (34.7),
caffeine (36.2), and MSG (37.2) were perceived not to be signicantly different from each other. The aroma only controls tested
amongst samples containing lactic acid (42.1) were perceived to
be signicantly more intense than controls presented amongst
NaCl and sucrose. These differences clearly demonstrated a contrast effect that depended on whether the taste type enhanced or
suppressed cheese avour intensity (Lawless & Heymann, 2010).

3.2. Effect of aroma level on cheese avour intensity

Based on mean panel ratings, cheese avour intensity was signicantly (p < 0.001) enhanced by aroma level for each single taste
type, and for the mixture of the ve tastes. As expected, cheese avour intensity signicantly increased as aroma level increased for
all taste types. Further effects were found when the differences
in cheese avour intensities for taste containing samples were
compared with the controls containing aroma only (Fig. 4).
Adding sucrose enhanced cheese avour intensity at all aroma
levels compared with the aroma only control (Fig. 4a). The
enhancement in avour intensity, however, was not signicantly
different between aroma levels.
Furthermore, the magnitude of enhancement of cheese avour
intensity by NaCl was signicantly higher (p < 0.01) at high aroma
level than it was at both low and medium aroma levels (Fig. 4b).
Addition of MSG enhanced the cheese avour intensity at both
low and medium aroma levels, but caused suppression at the high
aroma level (Fig. 4c). The cheese avour intensity at low and medium aroma levels was signicantly higher (p < 0.001) than the avour intensity at high aroma level. This was the source of the
signicant MSG concentration  aroma concentration interaction
as described in Section 3.1.

20

(a)

Difference in cheese flavour

intensity

10
5
0
-5
-10
-15
-20
Low

Medium
Aroma

(c)
b

0
-5
-10
-15
-20
Medium
Aroma

(e)

15
10
5

0
-5
-10
-15
-20
Low

Medium
Aroma

10

High

(b)

a
a

5
0
-5
-10
-15
-20
Medium
Aroma

High

(d)

20
15
10
5

c
a

Low

Medium
Aroma

0
-5
-10
-15
-20

High

20

15

Low

15
10

20

High

20

Low

Difference in cheese flavour

intensity

Difference in cheese flavour

intensity

Difference in cheese flavour

intensity

15

Difference in cheese flavour

intensity

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

Difference in cheese flavour

intensity

20

High

(f)
a

15
10
5
0
-5
-10
-15
-20
Low

Medium
Aroma

High

Fig. 4. Mean difference in cheese avour intensity from a taste-free control ( standard error) by aroma level when combined with (a) sucrose, (b) NaCl, (c) MSG, (d) lactic
acid, (e) caffeine, and (f) mixture of ve tastes. Means with the same superscripts are not signicantly different according to Fishers LSD.

Added lactic acid suppressed cheese avour intensity at both

low and medium aroma levels but had no effect at the high aroma
level (Fig. 4d). The differences in relative suppression were significant (p < 0.001). The avour intensity was suppressed most at low
aroma level.
Added caffeine suppressed cheese avour intensity at all aroma
levels to a small magnitude (Fig. 4e). There were no differences in
relative suppression. Caffeine therefore had minimal inuence on
the intensity of cheese avour.
The mixture of ve tastes enhanced cheese avour intensity all
aroma levels (Fig. 4f). Although a trend was apparent, there was no
signicant difference in relative enhancement.
3.3. Effect of tastes on taste intensity
In addition to cheese avour intensity, taste intensity was measured for each taste type. As expected, an increase in concentration
of a single taste signicantly increased the intensity of its corresponding taste attribute (all p < 0.001). The concentration ranges
of single tastes covered a wide range on the taste intensity scale.
Single tastes of sucrose, NaCl, and MSG signicantly increased
sweetness, saltiness, and umami, respectively, at each increasing
concentration level. The sourness of lactic acid and bitterness of
caffeine signicantly increased up to 0.28% lactic acid and 0.13%
caffeine, respectively.
All taste intensities signicantly (all p < 0.001) increased with
increasing concentrations of the mixture of ve tastes. Sweetness
signicantly increased up to taste level 4 whereas saltiness, umami, sourness, and bitterness signicantly increased up to taste level
5. The sweetness, saltiness, sourness, and bitterness ratings overall
were rated lower in the mixture compared with single tastes. The
sweetness of sucrose was signicantly suppressed by the other

tastes at level 4% (p < 0.01) and above (both p < 0.001) (sucrose at
4.78%, NaCl at 0.32%, MSG at 0.18%, lactic acid at 0.18%, and caffeine at 0.08%). The saltiness was signicantly enhanced at 0.13%
NaCl (p < 0.001) by 1.9% sucrose, 0.07% MSG, 0.07% lactic acid,
and 0.03% caffeine, and 0.2% NaCl (p < 0.05) by 3.01% sucrose,
0.11% MSG, 0.11% lactic acid, and 0.05% caffeine. Saltiness was signicantly suppressed at 0.5% NaCl by 7.57% sucrose, 0.28% MSG,
0.28% lactic acid, and 0.13% caffeine, and 0.8% NaCl by 12% sucrose,
0.45% MSG, 0.45%, lactic acid, and 0.2% caffeine (both p < 0.001).
The umami of MSG was not signicantly suppressed by the other
tastes. The sourness of lactic acid was signicantly suppressed by
equi-intense tastes at all lactic acid concentrations (all p < 0.001).
The bitterness of caffeine was signicantly suppressed by equi-intense tastes at all caffeine concentrations (all p < 0.001). These suppressions can be attributed to mixture suppression (Bartoshuk,
1975; Green, Lim, Osterhoff, Blacher, & Nachtigal, 2010). Based
on the comparison of taste intensities at the highest taste levels between the mixture and single tastes, the umami intensity did not
appear to be suppressed (difference of 0.3 points) to the same extent as sweetness, saltiness, sourness, and bitterness (difference of
32.7, 21.1, 49.3, and 32.1 points, respectively). The mutual suppression effect of sweetness, saltiness, sourness, and bitterness was
consistent with effects reported by Bartoshuk (1975). The relative
stability of umami ratings in the mixture does not appear to have
been previously reported.
3.4. Effect of aroma on taste intensity
Aroma level did not signicantly change the sweetness, saltiness, or sourness intensities of sucrose, NaCl, or lactic acid, respectively even though the aroma levels signicantly inuenced the
cheese avour intensity in each case as reported in Section 3.2.

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

(a)

60

Umami intensity

b
b

50
a

40

30
Low

Medium

High

Aroma level
60

(b)

Umami intensity

b
50

40

30
Low

Medium
Aroma level

High

(c)

60

Bitterness intensity

b
a

Low

Medium
Aroma level

50

40

30
High

Fig. 5. Mean taste intensities ( standard error) at three different aroma levels.
Taste intensities shown were for (a) umami of MSG, (b) umami of mixture of ve
tastes, (c) bitterness of caffeine. Means with the same superscripts are not
signicantly different according to Fishers LSD.

No signicant interactions of aroma concentration  sucrose concentration, aroma concentration  NaCl concentration, and aroma
concentration  lactic acid concentration were detected. This
meant that increasing concentrations of sucrose, NaCl, or lactic
acid increased taste intensity similarly across all aroma levels. A
lack of enhancement of taste intensity by aroma in the current
results, contradicts previous reports, particularly of saltiness
enhancement by aroma in cheese. However, in this study the variable was aroma level, whereas other studies have compared added
aroma with a no aroma condition (Lawrence et al., 2009, 2011).
These authors concluded that aroma could be used to compensate
for a reduced salt content by enhancing saltiness. In the current
study the lack of a signicant saltiness enhancement between aroma levels questions the ability of aroma to compensate for taste
reduction. It is also the case that use of an added aroma of different
character will result in a changed cheese avour character that
may not be acceptable.
Umami taste was signicantly (p = 0.01) enhanced by aroma
(Fig. 5a). Both medium and high aroma levels had more intense
umami taste than the low aroma level. A signicant (p < 0.05)
interaction of aroma concentration  MSG concentration was
detected, where the increase in umami intensity with increase in
MSG concentration was larger at low aroma concentration

compared to medium and high aroma concentrations. Aroma also

signicantly (p < 0.01) enhanced umami taste within the mixture
of ve tastes (Fig. 5b). The enhancement of umami from low to
high aroma for MSG singly and the mixture of ve tastes was 7.2
and 3.8 points, respectively, which was relatively small. In the case
of a mixture of ve tastants there was a signicant aroma concentration  taste concentration interaction. As the umami intensity
was enhanced by high aroma at tastant level 1 and 2, the overall
magnitude of increase in umami intensity between MSG levels at
high aroma levels were relatively smaller than at low and medium
aroma levels.
While the enhancement of umami taste by cheese aroma has
not previously been reported, umami is known to be an important
component of cheeses. Glutamate was found to be largely responsible for the umami taste in Cheddar and Swiss cheeses (Drake
et al., 2007). In Cheddar and Gouda cheese extracts, omitting umami tasting compounds increased intensities of specic attributes
such as bitterness, and reduced saltiness and also the overall intensity (Andersen et al., 2010; Drake et al., 2007; Toelstede & Hofmann, 2008a). Umami enhancement by aroma may present the
opportunity of using umami tastes as avour enhancers at lower
levels in concert with aroma, to achieve the same effect as using
umami tastes alone at higher levels.
The bitterness of caffeine was signicantly (p < 0.05) enhanced
by cheese aroma (Fig. 5c) when aroma level was increased from
either the low or medium level to the high aroma level. No significant interaction of aroma concentration  caffeine concentration
was detected, where the bitterness intensity with increasing caffeine concentrations was similar across all aroma concentrations.
However, similar to the result reported for umami, the magnitude
of bitterness enhancement was relatively small (enhancement of
4.49 points from low to high aroma). Changes in perceived bitterness due to cheese aroma have not been previously reported; bitterness is considered a defect and research has generally focused
on reducing this taste. The effects of aroma on perceived taste
intensity were stable across replicates, in agreement with those
found for perceived cheese avour intensity. These effects will be
further discussed in Section 3.5.
3.5. Possible mechanisms of cross-modal sensory interactions
Flavour perception, both taste and aroma perceived in concert,
has been proposed as being dependent on learning through experience over time (Small et al., 2007). Multiple exposures to a
tastearoma combination can promote implicit learning of the
association, which can later induce cross-modal sensory interactions (Stevenson, Boakes, & Prescott, 1998). Associations formed
as a result of learning between aroma and taste subsequently are
labelled as congruent combinations (Prescott, 2012), where congruency is the degree at which stimuli are perceptually in harmony
with each other (Schifferstein & Verlegh, 1996). This has been demonstrated with measurements of brain activity with functional
magnetic resonance imaging. The active areas in the brain specic
to gustation and olfaction appear to overlap with each other and
concurrent perception of matching taste and aroma results in supra-additive activity compared with taste or aroma perceived
alone (Eldeghaidy et al., 2011; Small, 2008). These types of signals
appear to only occur with familiar combinations of taste and aroma
and also have been speculated to translate to the intensity of avour perception. Perception of mismatched mixtures of taste and
aroma on the other hand has the opposite effect; the specic area
that previously showed supra-additive activity decreases. This effect was reported when vanilla aroma was combined with salt in
solution (Small et al., 2004). Mismatching taste and aroma have
also been shown to cause suppression in avour intensity (Stevenson et al., 1998). From this perspective the enhancement of cheese

J. Niimi et al. / Food Quality and Preference 31 (2014) 19

avour intensity by NaCl, sucrose, and MSG in the current study

seemed to be related to congruency due to familiarity of the combination of the tastes and aroma.
Another mechanism may also have been responsible for avour
enhancements, in particular with sucrose, due to its innate ability
to enhance avour intensity or even to have associations that were
learned early in life. This is consistent with the hypothesis of Green
et al. (2012) and Lim and Johnson (2011), where nutritive tastes
such as sweet, salty, and umami enhance retronasal aroma. To
add to this effect, Prescott, Johnstone, and Francis (2004) showed
that an association between an aroma with sweetness was induced
through learning by a single exposure. These authors demonstrated that exposure had increased smelled sweetness of the
odour whilst there was no change by the unexposed group. This
may mean that the ability of sucrose to enhance avour is not limited by the degree of familiarity of sweetness and aroma combinations. It may be, due to human nature to seek food with caloric
value that are sweet or salty tasting foods to form associations
with unfamiliar aromas relatively quickly, simply because many
foods have characteristic tastes of sweet, salty, or contain a combination of both.
Cross-modal sensory interaction studies in the scientic literature have reported effects with both nave and experienced assessors. Typically nave assessors are used as they tend to naturally
use a holistic and synthetic processing during avour perception
that facilitates the detection of interactions (Batenburg & van der
Velden, 2011; Hort & Hollowood, 2004). A synthetic approach
helps to integrate the perception cognitively. There are, however,
numerous caveats with using nave assessors concerning stability
of interactions. This may include lack of understanding of specic
attributes (Stone & Sidel, 2004) and the inuence of expectation
bias (Cardello, 1995), resulting in variable interaction effects across
replicates for some aroma-taste combinations (Green et al., 2012).
To add to this list, repeated exposure to an aromataste combination can increase assessors familiarity, resulting in a changed avour intensity compared with initial exposure (Prescott et al.,
2004). In using experienced assessors, there are advantages and
disadvantages also. Experienced assessors have the advantage of
acuity, ability to discriminate intensities, and repeatability in their
ratings. They have been shown to eliminate sensory interactions
however, due to their overly analytical evaluation approach (van
der Klaauw & Frank, 1996).
The main reason for using experienced assessors in the current
study was that the interest was in true or robust interactions,
rather than interactions that are biased by expectation of unfamiliarity with the mixed stimulus. The general lack of enhancement of
taste by aroma that was found may be partly attributed to the
experience of assessors used for the evaluations who adopted an
analytical approach to rating intensity of avour and taste separately. However, it may also be the case that cheese aroma cannot
enhance tastes such as saltiness and sweetness. On the other hand,
taste had a large and robust impact on cheese avour intensity, signicantly enhancing the contribution of aroma. This suggested that
avour is holistic, and not susceptible to an analytical dissection.
Rather than measuring taste and aroma independently when
mixed, it appears preferable to measure avour to detect stable
cross-modal sensory interactions.
The stability of cross-modal sensory interactions is important as
successful food products are repeatedly consumed. Steps taken in
the current study to facilitate the detection of interactions included
the provision of detailed instructions, providing as opposed to generating attributes by the panel to prevent excessive dissection, and
making certain that assessors understood and could recognise
attributes that were rated. The strong interaction effect on cheese
avour intensity appeared to be repeatable, which was shown with
the overall non-signicant replicate effects. Assessors were

exposed to a specic taste and aroma combination 30 times in total

(ve taste concentrations, three aroma levels, repeated).
4. Conclusions
Combinations of tastes and aroma can either enhance or suppress cheese avour. Sucrose, NaCl, and a mixture of tastes enhanced cheese avour intensity at all aroma levels. MSG also
enhanced cheese avour intensity, but was dependent on aroma
level. Lactic acid, unlike the other tastes, suppressed cheese avour
intensity, particularly at high acid levels, and at low and medium
aroma levels. A mixture of ve tastes enhanced cheese avour
intensity more than any of the single tastes. The magnitude of
cheese avour intensity change was not linearly dependent on
taste concentrations, and above a certain concentration the addition of a higher level of a taste did not result in further enhancement or suppression. Umami and bitterness intensities were
enhanced by cheese aroma.
Acknowledgements
The donation of pure food grade aroma compounds by Givaudan was greatly appreciated. J.N. also acknowledges Fonterra Cooperative Group and the Foundation for Research, Science and
Technology (New Zealand) for providing funding for this project.
J.N. thanks the members of CSIRO Food and Nutritional Sciences
for their help and assistance.
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