Acta Psychologica 132 (2009) 190200

Contents lists available at ScienceDirect

Acta Psychologica
journal homepage: www.elsevier.com/locate/actpsy

An fMRI investigation on image generation in different sensory modalities:

The inuence of vividness
M. Olivetti Belardinelli a,b,*, M. Palmiero a,b,c, C. Sestieri d,e, D. Nardo a,b, R. Di Matteo b,d, A. Londei b,
A. DAusilio b, A. Ferretti d,e, C. Del Gratta d,e, G.L. Romani d,e
a

Department of Psychology, Sapienza University of Rome, Italy

ECONA, Interuniversity Centre for Research on Cognitive Processing in Natural and Articial Systems, Italy
Perceptual and Dynamic Laboratory, Riken Brain Science Institute, Japan
d
Department of Clinical Sciences and Bio-Imaging, G. dAnnunzio University, Chieti, Italy
e
ITAB, Institute for Advanced Biomedical Technologies, G. d Annunzio University Foundation, Chieti, Italy
b
c

a r t i c l e

i n f o

Article history:
Received 22 July 2008
Received in revised form 14 June 2009
Accepted 22 June 2009
Available online 19 August 2009
PsycINFO classication:
2300
2340
Keywords:
Imagery
Vividness
Sensory-modality
Representational format
Simulation

a b s t r a c t
In the present fMRI study the issue of the specic cortices activation during imagery generation in different sensory modalities is addressed. In particular, we tested whether the vividness variability of imagery
was reected in the BOLD signal within specic sensory cortices. Subjects were asked to generate a mental image for each auditory presented sentence. Each imagery modality was contrasted with an abstract
sentence condition. In addition, subjects were asked to ll the Italian version of the Questionnaire Upon
Mental Imagery (QMI) prior to each neuroimaging session. In general, greater involvement of sensory
specic cortices in high-vivid versus low-vivid subjects was found for visual (occipital), gustatory (anterior insula), kinaesthetic (pre-motor), and tactile and for somatic (post-central parietal) imagery modalities. These results support the hypothesis that vividness is related to image format: high-vivid subjects
would create more analogical representations relying on the same specic neural substrates active during
perception with respect to low-vivid subjects. Results are also discussed according to the simulation
perspective.
2009 Elsevier B.V. All rights reserved.

1. Introduction
Mental imagery is considered to be an important component of
conscious experience (Paivio, 1969), although it is held to usually
occur with a lower intensity when compared to real perception
(Fallgatter, Mueller, & Strik, 1997), since it is generated when perceptual information is retrieved from memory, and people are seeing with their minds eye (Kosslyn, 1994; Marks, 1973), or
hearing with their minds ear (Halpern, 1988; Pitt & Crowder,
1992). In recent years, additional experimental evidence demonstrated that people are able to generate mental images also in tactile (Yoo, Freeman, McCarthy, & Jolesz, 2003), kinaesthetic
(Jeannerod, 1995), olfactory (Djordjevic, Zatorre, Petrides, &
Jones-Gotman, 2004; Elmes, 1998), and gustatory (Kikuchi, Kubota,
Nisijima, Washiya, & Kato, 2005) modalities.
Although many researchers argue that mental images in different sensory modalities preserve key elements of perceptual stimuli
* Corresponding author. Address: Department of Psychology, Sapienza University of Rome, and ECONA, Italy. Tel.: +39 06 49917609; fax: +39 06 4462449.
E-mail address: marta.olivetti@uniroma1.it (M. Olivetti Belardinelli).
0001-6918/$ - see front matter 2009 Elsevier B.V. All rights reserved.
doi:10.1016/j.actpsy.2009.06.009

(Algom & Cain, 1991; Carrasco & Ridout, 1993; Decety & Michel,
1989; Halpern, 1988; Intons-Peterson, 1992; Kosslyn, 1980; Kosslyn, Ball, & Reiser, 1978; Parsons, 1994; Shepard & Metzler, 1971),
others state that these representations rely on abstract symbols of
the sort used in language (Anderson & Bower, 1973; Pylyshyn,
1973, 1979, 1981, 2002). This debate leads to the issue whether
mental imagery and perception share the same mechanisms. The
advent of neuroimaging techniques allowed researchers to shed
light on the issues of modality-specicity and imagery format
(Farah, 2000), addressing at least two different questions: (a) Is
there an anatomical separation between the cortical areas serving
imagery and those serving perception? (b) Are the areas used for
imagery a subset of those engaged in perception?
Contrasting results have been provided so far. On one hand,
some studies did not report the activation of early sensory areas
during imagery tasks (Bunzeck, Wuestenberg, Lutz, Heinze, &
Jancke, 2005; DEsposito et al., 1997; Ishai, Ungerleider, & Haxby,
2000a; Jahn et al., 2004; Mellet, Tzourio, Denis, & Mazoyer, 1998;
Mellet et al., 2000). On the other hand, other studies found that
perception and imagery share neural networks involving early sensory areas during visual (Amedi, Malach, & Pascual-Leone, 2005;

M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

Cui, Jeter, Yang, Montague, & Eagleman, 2007; Kosslyn, Thompson,

Kim, & Alpert, 1995; Kosslyn et al., 1999), auditory (Bunzeck et al.,
2005; Hoshiyama, Gunji, & Kakigi, 2001), tactile (Yoo et al., 2003),
olfactory (Bensa, Sobel, & Khan, 2007; Djordjevic, Zatorre,
Petrides, Boyle, & Jones-Gotman, 2005), gustatory (Kikuchi et al.,
2005; Kobayashi et al., 2004), and kinaesthetic (Dechent, Merboldt,
& Frahm, 2004; Lotze et al., 1999; Porro et al., 1996; Roth et al.,
1996; Szameitat, Shen, & Sterr, 2007) imagery. Interestingly, a previous fMRI study from our group (Olivetti Belardinelli et al., 2004a,
2004b) investigated in the same experiment imagery for seven different sensory modalities (visual, auditory, kinaesthetic, olfactory,
gustatory, tactile and somatic) triggered by visually presented sentences. Results revealed that all imagery modalities activated
mainly two higher associative areas, located in the fusiform gyrus
(BA 37) and in the inferior parietal lobule (BA 40), with a slightly
different pattern across modalities.
Although these divergent results may be partly explained by
considering small task differences (e.g., types of images, object categories, stimulus complexity, task requirements and modality of
administration; Lambert, Sampaio, Scheiber, & Mauss, 2002), a major confound may be represented by individual differences in the
ability to generate mental images. Traditionally, attempts to relate
individual differences in imagery abilities were made using self-report questionnaires (Betts, 1909; Galton, 1880). Despite the general methodological criticisms for such methods, questionnaires
were developed to test if vividness represents a general imagery
ability (Sheehan, 1967) or a modality-specic characteristic (Switras, 1978; White, Ashton, & Law, 1974). Furthermore, specic
questionnaires were developed for visual (Marks, 1973), motor
(Isaac, Marks, & Russel, 1986), auditory (Gissurarson, 1992) and
olfactory (Gilbert, Crouch, & Kemp, 1998) imagery modalities.
In recent years neuroimaging techniques addressed the issue of
the relationship between subjective and objective measures of
imagery. For instance, some studies showed that subjective measures of vividness in visual imagery were associated with BOLD
changes in the visual cortex (Amedi et al., 2005; Cui et al., 2007).
In the present research, thus, we faced the question whether individual variability in imagery vividness, assessed by means of the
Italian version on the Questionnaire Upon Mental Imagery, may
be reected in the activity of the corresponding early sensory cortex for each modality. We predicted that greater involvement of
modality-specic cortices should be associated with higher subjective measures of vividness in each sensory modality, representing
the specic sensory properties of vivid imagery.

191

(e.g., the action of writing); another one related to internal proprioceptive sensations: Somatic (e.g., the sensation of fatigue);
and the last one related to Abstract concepts (e.g., racial prejudice).
The experimental sentences were in part derived from a previous
study (Olivetti Belardinelli et al., 2004a, 2004b) and in part newly
constructed on the basis of the same criterion used there (for an
English version of the complete list of sentences, see Appendix A).
Category distinctiveness of the new sets of experimental sentences was tested. Thirty one subjects (different from those who
participated in the fMRI study) lled in a questionnaire and rated
the sentences along two dimensions: the dominant imagery
modality (multiple choice, including also the abstract option)
and the level of vividness (7-point Likert scale). Participants responses signicantly matched the item classication (p < 0.001),
assigning every item to the correct category; moreover modalityspecic items rated as more vivid than abstract items (modalityspecic items: mean 5.15, SD 1.48; abstract items: mean 2.13, SD
1.76; t = 14.74, p = 0.000). The ANOVA on mean vividness scores
contrasting category (eight levels seven imagery modality plus
abstract condition) and classication (two levels matching and
no-matching)
revealed
a
main
effect
for
category
(F[1,30] = 108,2411, p < 0.001), a main effect for classication
(F[7,210] = 21,5074, p < 0.001) and a signicant interaction between them (F[7,210] = 5,7524, p < 0.001). The result indicates that
coherently classied items were judged more vivid than nonmatching items for all categories except for abstract items; moreover within the coherently classied group only the abstract items
differed signicantly in vividness scores from all the other categories (Tukey p < 0.001).
The selected sentences were rst digitally recorded and then
post-processed. The gender connotation of the speakers voice
was concealed using a sound-editing software, in order to reduce
the BOLD activation due to the recognition of the speakers gender
(Imaizumi et al., 2004). A further pre-experimental test, with ten
women and ten men, ascertained that each subject attributed the
speakers voice to a person of her/his own sex.
In the fMRI session, the auditory stimuli were delivered through
a pneumatic headset, designed to minimize interference from the
scanner noise. Sound sampling rate was 22,000 Hz with 16-bit resolution. Stimuli were presented in stereo, at a sound pressure level
(SPL) of 85 dB. Despite the high-frequency cut-off of the pneumatic
device (estimated around 4 kHz), subjects reported that they were
able to clearly hear and understand the meaning of the sentences.
2.3. Procedure

2. Materials and methods

2.1. Subjects
Nine, healthy right-handed university students (mean
age = 25.2, SD = 3.7) participated in this study. Only female subjects were selected to reduce the gender-related variability that
has been reported in imagery tasks (White, Ashton, & Braown,
1977). All of them had normal hearing and vision. Participants gave
informed consent for a protocol approved by the local Institutional
Ethics Committee and were paid for their participation.
2.2. Stimuli
Subjects were stimulated with a set of 96 (12 for each category)
auditory short sentences belonging to eight categories: ve categories related to different sensory modalities (e.g., Visual: to see a
candle; Auditory: to hear a shot; Tactile: to touch something
hard; Olfactory: the smell of alcohol; Gustatory: the salty
taste); one category related to self-perceived motion: Kinaesthetic

Prior to the scanning session, participants were asked to complete the Italian versions of the Edinburgh Handedness Inventory
(Oldeld, 1971) to assess their handedness, and the Questionnaire
of Mental Imagery (Sacco & Reda, 1998) to assess their vividness in
mental imagery generation in different sensory modalities. The
subsequent fMRI session lasted for approximately 40 min, during
which both functional and anatomical data were acquired. The paradigm was a classic block paradigm, divided in three runs. Each run
consisted of eight blocks (one of imagery in each of the seven sensory modalities plus one abstract block) lasting 28 s and intermixed with rest periods of 20 s. Therefore each run lasted
7 min. During rest periods, subjects were instructed to maintain
xation on a black cross presented at the centre of a white screen.
In each block, a cycle of four auditory sentences belonging to one
modality at a time was presented. Each cycle consisted of one sentence, lasting 2.5 s, followed by 4.5 s for mental images generation.
Subjects were instructed to listen to each sentence, create a mental
image of its content and maintain that image until the next sentence was delivered. For the whole fMRI session subjects were
explicitly instructed not to move at all, in order to avoid artefacts.

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

2.4. Functional imaging

Data were acquired with a Siemens Magnetom Vision 1.5 T
scanner, by means of T2*-weighted echo planar imaging (EPI) free
induction decay (FID) sequences with the following parameters: TR
2698 ms, TE 60 ms, matrix size 64  64, FOV 256 mm, in-plane
voxel size 4  4 mm, ip angle 90, slice thickness 5 mm and no
gap. In each run, a total of 155 functional volumes were acquired,
consisting of 25 trans-axial slices, including the whole brain. A
high-resolution structural volume was acquired at the end of the
session via a 3D MPRAGE sequence with the following features: axial, matrix 256  256, FoV 256 mm, slice thickness 1 mm, no gap,
in-plane voxel size 1  1 mm, ip angle 12, TR = 9.7 ms, and
TE = 4 ms.
3. Data analysis

nally images were smoothed with a 12 mm full-width-at-halfmaximum (FWHM) isotropic Gaussian kernel to increase the signal-to-noise-ratio and allow a group analysis.
3.2. Statistical analysis on vividness scores
Vividness ratings obtained using the Italian version of the
Questionnaire of Mental Imagery (QMI) (Sacco & Reda, 1998) were
analysed averaging the scores that subjects attributed to respective
items for each imagery modality. Therefore, each subject got seven
vividness imagery scores, (Table 1-A). In order to split between
high and low-vivid subjects a k-means overall analysis was
performed on these scores, resulting in two groups with their
respective averages in each imagery modality (Table 1-B). Finally,
considering the Euclidean distances, a high-vivid group (four
subjects) and a low-vivid group (ve subjects) were formed
(Table 1-C).

3.1. Image preprocessing

3.3. fMRI statistical analysis
Data were analysed using MATLAB 7.0 and SPM2 (http://
www.l.ion.ucl.ac.uk/spm). After discarding the rst ve images
of each session to suppress T1 saturation effects, the 465 whole
brain volume images (155 for each session) were realigned and
resliced to correct for interscan head movements and then corrected for differences in acquisition time between slices during
sequential imaging. The anatomical image of each subject was
co-registered to the functional mean image calculated during the
realignment. Anatomical images were then spatially normalized
to standard brain space (MNI T1 template) using a nonlinear spatial transformation with a 7  8  7 basis function. The estimated
nonlinear transformations were then used to reslice the corresponding functional images with a voxel size of 4  4  5 mm. Fi-

Statistical analyses were performed on a voxel-by-voxel basis

using the General Linear Model and the theory of Gaussian Random Fields as implemented in SPM2 on the 465 preprocessed
images. The design matrix was generated using a block design on
a canonical HRF basis function. High pass ltering with a cut-off
period of 128 s was carried out and auto-regression was used to
correct for serial correlations. The rest condition was modelled
implicitly in the design and served as a baseline. At the single-subject level, eight experimental conditions were modelled as
explanatory variables (seven imagery modalities visual, auditory,
tactile, gustatory, olfactory, motor, and somatic plus the abstract
concepts) and a voxel-wise t statistic for each condition was

Table 1
(A) Vividness scores obtained on the Vividness Imagery Questionnaire (QMI Italian Version): low scores indicate high-vividness ratings; high scores indicate low-vividness
ratings. (B) Average of each group after K-means analysis, and relative t-test: group 1, low-vividness; group 2, high-vividness. (C) Group members and Euclidean distance from
each respective group.
A
S1
S2
S3
S4
S5
S6
S7
S8
S9

Visual

Auditory

Tactile

Olfactory

Gustatory

Kinaesthetic

Somatic

1.4
1.8
2.4
2.4
1.6
1.6
1.2
2
2.4

3
2
1.4
1.4
1.2
1
2
1.8
2

2
1.6
1.2
2.4
1.6
2.4
1.2
1.4
2

2.2
1.8
1.2
1.6
3
1.8
1.2
2.8
2.2

2
1.4
1.2
1.8
2.2
1.2
1.6
1.4
2.2

2.6
2
1.6
3
2.6
1.4
1.4
2.6
3.4

2
1.6
1.2
2.8
2
2
1.4
2.2
2.4

B
Group 1

Group 2

Visual
Auditory
Tactile
Olfactory
Gustatory
Kinaesthetic
Proprioceptive

1.960000
1.880000
1.880000
2.360000
1.920000
2.840000
2.280000

1.750000
1.600000
1.600000
1.500000
1.350000
1.600000
1.550000

t-Test

0.000657145

C
High-vivid subjects
S2
0.244219

S3
0.35

S6
0.443605

S7
0.345895

Low-vivid subjects
S1
0.500057

S4
0.475455

S5
0.430017

S8
0.330886

S9
0.305754

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

Table 2
List of locations of the regions of interest referred to previous related studies.
Reference

Anatomical area

BA

Coordinates

Kan et al. (2003)

Kosslyn et al. (1999)

Left Fusiform Gyrus

Occipital Cortex

Bunzeck et al. (2005)

Heschl Gyrus

37
17
(18/19)
(41/42)

Djordjevic et al. (2005)

Kikuchi et al. (2005)
Yoo et al. (2003)
Szameitat et al. (2007)

PiriformOrbitoFrontal Cortex
Anterior Insula
Post-Central Gyrus
Pre-Central Gyrus

11
13
(1/2/3)
4
6

calculated against the rest condition, resulting in a series of statistical parametric maps (SPMs). The contrast images calculated for
each voxel at the single-subject level were then entered into the
group random-effect analysis.
In the group analysis, the different conditions, estimated at the
single-subject level, were contrasted as t statistics (random-effect)
comparing each imagery modality with the abstract condition.
Activated regions were identied as signicant only if they reached
a height threshold of p < 0.05, corrected (FDR) for multiple comparisons at voxel-level, with a cluster size equal or exceeding 10 contiguous activated voxels (except for post-central activation in the
somatic modality in the two-sample t-test analysis). MNI coordinates of the local maxima were then transformed into Talairach
coordinates (Talairach & Tournoux, 1988) using the MNI2Tal provided by M. Brett (http://www.mrc-cbu.cam.ac.uk/imaging/common/mnispace.shtml) and activated areas were labelled
according to the Talairach Daemon Database (http://ric.uthscsa.e-

Y
45
2
34
44
40
27
44
57
18
32

Z
53
88
86
10
20
6
10
27
30
7

6
12
12
8
8
11
2
38
60
50

du/resources/body.html). In addition, a small volume correction

(SVC) was performed on both hemispheres using a 10 mm radius
within anatomical regions of interest (ROIs). The locations of the
regions of interest were based on a priori hypotheses drawn from
previous related studies (see Table 2).

4. Results
4.1. Relevant activation for the whole sample
The one sample t-test analysis comparing each imagery modality versus the abstract condition was performed to explore what
process imagery modalities have in common once critical components involved both in imagery and in the abstract condition are
subtracted (e.g., task initiation and maintenance, attention, and
working memory). In general, the activation of the left fusiform

Table 3
Stereotaxic coordinates (Talairach & Tournoux, 1988), anatomical locations, and T scores of (local) peak activations for the comparison Imagery Modality > Abstract. The X sign
indicates Small Volume Correction applying.
Imagery modality

Antomical area

BA

Coordinates
X

Cluster
Y

SVC

p (cor)

Visual

L Fusform Gyrus
L Fusform Gyrus

37
37

48
40

64
58

13
6

0.035

113

4.74
5.01

Auditory

L Fusiform Gyrus
L Middle Temporal Gyrus
L Caudate Tail

37
37

45
55
36

58
58
35

7
1
2

0.049

14

3.52
3.52
4.54

Tactile

L Superior Temporal Gyrus

L Fusform Gyrus
L Caudate Tail

22
37

44
40
36

27
47
39

2
8
2

0.032
0.032

14
14

2.60
5.27
2.70

L
L
L
L
L
L

7
7
7
30
37

16
20
16
16
48
32

71
64
60
39
64
42

50
40
49
7
11
7

0.609

85

0.039
0.039

16
16

4.43
2.93
2.89
3.44
5.04
3.41

Olfactory

Precuneus
Precuneus
Precuneus
Parahippocampal Gyrus
Fusform Gyrus
Hippocampus

Gustatory

L Fusform Gyrus
L Caudate Tail

37

51
36

59
27

10
3

0.049

10

4.38
5.25

Kinaesthetic

L
L
L
L
L
L
L
L

2
5
6
6
20
37
40

24
36
20
16
32
24
44
24

36
44
17
13
36
40
40
56

66
62
56
47
15
15
57
31

0.339

64

0.009

43

0.027
0.042
0.339
0.042

22
15
64
15

4.09
4.12
9.31
5.07
3.82
3.46
4.42
5.02

19
19
36
37
37

40
40
28
51
40

75
79
32
63
47

13
17
15
14
14

0.000
0.001
0.000
0.001
0.000

89
53
89
53
89

9.40
7.00
5.79
9.09
7.56

Somatic

Post-Central Gyrus
Paracentral Gyrus
Middle Frontal Gyrus
Medial Frontal Gyrus
Fusform Gyrus
Fusform Gyrus
Inferior Parietal Lobule
Cerebellum

R Fusiform Gyrus
L Fusform Gyrus
R Limbic Parahippocampal Gyrus
L Fusform Gyrus
R Fusiform Gyrus

X
X

X
X
X

194

M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

gyrus (BA37) was found for all the imagery modalities. The somatic
imagery yielded also the activation of the right fusiform gyrus
(BA37) and of the bilateral occipital fusiform gyrus (BA19),
whereas, the kinaesthetic imagery resulted in the activation of
the left fusiform gyrus (BA20). Activations are listed in Table 3
and shown in Fig. 1.
4.2. Modality-specic activation
The two-sample t-test analysis comparing high-vivid versus
low-vivid subjects, within each imagery modality contrasted with
the abstract condition, showed signicantly greater activations in
sensory specic areas for high-vivid compared to low-vivid subjects. Specically, we observed greater activity for the high-vivid

subjects in: the left inferior occipital gyrus (BA17/18) for visual
imagery; the left anterior insula (BA13) for gustatory imagery;
the right post-central gyrus (BA2) for tactile and somatic imagery;
and the left pre-central gyrus (BA4) for kinaesthetic imagery. Activations are listed in Table 4 and presented in Fig. 2.
In order to clarify the differences between the activation patterns of high- and low-vivid subjects, a ROI analysis (implemented
as SVC) was performed in relevant sensory specic areas. As shown
in Fig. 3, activity in these areas was enhanced in high-vivid subjects
and deactivated in low-vivid subjects. This was observed for all
sensory modalities we tested. In general, the analysis showed signicant activities in each ROI only during the associated sensory
modality (i.e. visual cortex ROI showed activity during visual imagery only).

Fig. 1. Activation of the left Fusiform gyrus for the comparison Imagery Modality > Abstract.

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

Table 4
Stereotaxic coordinates (Talairach & Tournoux, 1988), anatomical locations, and T scores of (local) peak activations for the comparison Imagery Modality > Abstract in high-vivid
with respect to low-vivid subjects. The X sign indicates Small Volume Correction applying.
Imagery modality

Antomical area

BA

Coordinates
X

Cluster
Y

p (cor)

SVC
K

Visual

L Inferior Occipital Gyrus

L Inferior Occipital Gyrus

17
18

16
32

90
82

8
4

0.029
0.021

21
27

4.53
5.52

X
X

Auditory

L Middle Frontal Gyrus

L Middle Frontal Gyrus
L Middle Frontal Gyrus
R Transverse Temporal Gyrus
L Transverse Temporal Gyrus

10
10
10
42
42

44
28
36
67
56

47
63
59
19
16

11
11
11
10
14

0.940

90

0.194
0.145

23
47

4.83
3.76
3.62
5.80
4.86

X
X

Tactile

R Post-Central Gysrus
R Superior Frontal Gyrus
R Inferior Parietal Lobue
L Cerebellum (Anterior Lobe)

2
10
40

55
28
48
16

25
47
40
44

52
16
53
23

0.038
0.901
0.909
0.367

20
33
32
86

4.34
5.78
4.34
5.89

Olfactory

R Middle Frontal Gyrus

L Middle Frontal Gyrus
L Middle Frontal Gyrus
R Insula
L Fusiform Gyrus
R Inferior Parietal Lobule

6
10
10
13
37
40

48
32
32
40
48
48

14
50
40
11
64
40

45
7
16
6
11
53

0.035
0.288
0.288
0.075
0.039
0.975

19
536
26
7
16
148

5.07
5.48
3.60
3.10
5.47
4.34

L
L
L
L

Middle Frontal Gyrus

Anterior Insula
Superior Temporal Gyrus
Transverse Temporal Gyrus

10
13
38
42

44
36
32
56

47
16
3
16

11
5
13
14

0.948
0.021
0.936
0.145

90
28
95
47

4.83
3.82
2.95
4.86

L Post-Central Gyrus
L Pre-Central Gyrus
L Cerebellum (Anterior Lobe)
R Cerebellum (Anterior Lobe)

3
4

32
32
12
16

25
21
41
44

43
52
32
27

0.017
0.017
0.000
0.000

14
14
236
236

6.39
6.39
9.30
12.34

R Post-Central Gyrus
R Superior Parietal Lobule
R Superior Parietal Lobule

2
7
7

51
40
28

28
67
60

57
49
50

0.047

3.54
3.86
3.04

Gustatory

Kinaesthetic

Somatic

X
X

X
X
X

Fig. 2. Activation of sensory specic cortices for each imagery modality contrasted with abstract condition in high-vivid subjects with respect to low-vivid subjects.

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

Fig. 3. Histograms of contrast estimates and 90% Condential Interval for each ROI contrasting high- versus low-vividness for each imagery modality. For visual, auditory
modalities, the contrasts are shown in two different areas: left BA 17 and left BA 18 for visual imagery.

4.3. Correlation analysis

A regression analysis was also performed to regress the BOLD
signals onto the vividness scores and to better predict the activations in modality-specic areas. This analysis was done considering the contrasts at single-subject level between imagery
modality and abstract condition. In general, specic-modality cortices were found positively correlated with vividness scores for visual, tactile, gustatory, kinaesthetic and somatic imageries. In
particular, activations of the right lingual and inferior occipital
gyrus (BA18), as well as of the bilateral middle occipital gyrus
(BA19) were found for the visual imagery modality. In addition,
the right post-central gyrus (BA2) for tactile imagery, the left anterior insula (BA13) for gustatory imagery, the left pre-central gyrus
(BA6) for kinaesthetic imagery, and the right post-central gyrus
(BA3) for somatic imagery were found activated signicantly. For
the complete list of activations, refer Table 5.
5. Discussion
In the last decade, neuroimaging studies offered critical insights
into the long standing mental imagery debate. In fact, the neural
bases of mental image generation have been investigated in all
the sensory modalities, showing that both modality-specic and
a-specic areas are involved. Nevertheless, few studies focused
on the differences or similarities for all imagery modalities and

on the possible inuence of vividness variability on cortical activity. In this study we tested whether the fMRI BOLD signal was
modulated in sensory cortices according to the individual variability in vividness in all imagery modalities, using a set of sentences
similar to the one originally used by Olivetti Belardinelli et al.
(2004a, 2004b).
5.1. Relevant activations to all subjects (high- and low-vivid)
We compared each imagery modality with the abstract condition in the whole sample in order to better investigate the processes underlying the imagery operations, excluding attentional,
conceptual and working memory processes. The left fusiform gyrus
(BA37) was found active for all imagery modalities. In general, this
area has been related to semantic retrieval during a property verication task when associated false trials were present and in absence of explicit mental imagery instructions (Kan, Barsalou,
Solomon, Minor, & Thompson-Schill, 2003). Moreover, it has been
related to visual mental imagery (DEsposito et al., 1997), object
recognition (Ishai, Ungerleider, Martin, & Haxby, 2000b), and retrieval of object information (Price, Noppeney, & Phillips, 2003).
Nevertheless, since we used verbal cues to elicit images, the left
fusiform activation may also reect a visual semantic retrieval
(Thompson-Schill, Aguirre, DEsposito, & Farah, 1999), which gives
rise to multi-modal mental representation (e.g., visuo-olfactory, visuo-gustatory, etc., as shown in Olivetti Belardinelli et al., 2004a),

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

Table 5
Stereotaxic coordinates (Talairach & Tournoux, 1988), anatomical locations, and T scores of (local) peak activations for the comparison Imagery Modality > Abstract regressed
onto the vividness ratings. The X sign indicates Small Volume Correction applying.
Imagery modality

Antomical area

BA

Coordinates
X

Visual

Cluster
Y

p (cor)

SVC
K

R Middle Frontal Gyrus

R Middle Frontal Gyrus
L Middle Frontal Gyrus
R Superior Frontal Gyrus
L Superior Fronta Gyrus
R Superior Frontal Gyrus
R Lingual Occipital Gyrus
R Inferior Occipital Gyrus
R Middle Occipital Gyrus
L Middle Occipital Gyrus
L Posterior Cingulate
R Posterior Cingualte
R Cingualte Gyrus
R Anterior Cingualte
R Sovramarginal Gyrus
L Inferior Parietal Lobule

6
6
6
9
10
10
18
18
19
19
24
30
31
32
40
40

40
44
28
16
24
24
16
36
48
24
4
8
16
8
40
40

6
14
2
52
44
48
82
86
74
82
34
50
53
43
41
45

46
45
42
20
16
21
9
4
6
17
20
16
26
12
30
38

0.042
0.001
0.000
0.032
0.000
0.032
0.000
0.042
0.000
0.007
0.000

52
107
125
55
125
55
401
52
401
19
126

0.032
0.000

55
401

R Middle Frontal Gyrus

R Insula
L Lingual Gyrus
R Posterior Cingulate
R Angular Gyrus
R Superior Temporal Gyrus
L I nferior Frontal Gyrus
R Inferior Frontal Gyrus

10
13
19
23
39
42
44
47

36
40
28
4
45
67
51
51

51
16
74
30
68
19
1
39

3
13
1
20
31
10
18
2

0.77

102

0.000
0.000
0.000
0.078
0.131
0.085

1438
1438
1438
35
18
32

5.05
3.02
10.06
5.7
11.3
2
4.35
2.34

Tactile

R Post-Central Gyrus
R Anterior Insula
R Post-Central Gyrus
L Lingual Gyrus
R Superior Temporal Gyrus
L Post-Central Gyrus
R Cerebellum (Anterior Lobe)

2
13
5
18
38
43

48
36
32
12
51
51
28

28
26
44
82
19
18
52

52
15
57
9
14
19
27

0.005
0.001
0.005
0.000
0.001
0.000
0.035

27
146
27
669
146
387
74

7.4
4.02
5.99
6.65
7.02
12.3
4.35

Olfactory

R Middle Frontal Gyrus

R Middle Frontal Gyrus
R Middle Frontal Gyrus

10
10
10

45
28
40

50
55
52

11
20
20

0.996

93

3.43
3.38
3.37

Gustatory

L Anterior Insula
L Anterior Insula
L Post-Central Gyrus

13
13
43

40
36
59

1
7
6

14
14
19

0.05
0.05
0.05

16
16
16

4.27
4.21
4.4

Kinaesthetic

R Medial Frontal Gyrus

R Medial Frontal Gyrus
L Pre-Central Gyrus
L Middle Frontal Gyrus
R Cuneo
L Inferior Parietal Lobule
L Inferior Parietal Lobule

6
6
6
6
18
40
40

4
12
32
40
24
55
59

17
20
6
2
84
29
30

52
56
33
41
23
38
29

0.049

22

0.05

12

0.95

90

6.45
5.8
4.57
3.1
3.35
4.19
3.78

R Post-Central Gyrus
L Medial Frontal Gyrus
R Medial Frontal Gyrus
L Superior Temporal Gyrus
L Inferior Frontal Gyrus

3
6
6
22
44

24
4
12
63
59

28
9
24
50
16

57
47
56
12
13

0.002

545

0.004

494

Auditory

Somatic

as well as cross-modal interaction (Olivetti Belardinelli et al.,

2004c).
5.2. Modality-specic activations
When the abstract condition was subtracted from the imagery
generation process, activation of sensory specic cortices appears
to be modulated by vividness scores and critically more signicant
activity was found in high-vivid subjects only with respect to lowvivid, except for auditory and olfactory imagery. Regarding auditory imagery, the lack of signicant modality-specic activation
may be due to the interference of the scanner noise on the auditory
image formation process. In fact, contrasting an experimental and
baseline condition with continuous scanner background noise,
Gaab, Gabrieli, and Glover (2007) demonstrated that both may lead

4.03
7.13
6.06
4.92
4.9
4.77
6.95
7.5
4.74
5.05
9.27
4.27
4.36
4.89
8.36
8.28

X
X
X

6.5
7.56
7.21
5.25
5.25

to signal decreases, with a greater effect in Heschls gyrus. Regarding olfactory imagery, the lack of an olfactory activation leads to
the conclusion that the vividness scores related to olfactory imagery do not predict olfactory specic-modality activations, probably
depending on the difculty in generating vivid images of smells
(Lawless, 1997), especially when they are verbally cued (Herz,
2000).
With respect to conditions revealing signicant activity,
although correlation and group analyses do not completely overlap, both analyses revealed that modality-specic activations are
modulated by vividness level for visual, tactile, gustatory, kinaesthetic and somatic mental imageries.
In particular, for visual imagery, the activation of BA17 (group
analysis) or BA18 (both group and correlation analysis) was found.
Several studies provided no evidence of these activations for

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

instance, when participants generated images of objects, faces or

animals starting from concrete nouns or sentences (DEsposito
et al., 1997; Ishai et al., 2000a; Mellet et al., 1998; Olivetti Belardinelli et al., 2004a, 2004b), as well as when individuals were required to generate and inspect letters (Goldenberg et al., 1989;
Gulyas, 2001), or previously memorized maps (Ghaem et al.,
1997; Mellet, Tzourio, Denis, & Mazoyer, 1995; Mellet et al.,
2000). Nevertheless, early visual cortex activation was found in
many other studies (Goldenberg et al., 1989; Goldenberg, Steiner,
Podreka, & Deecke, 1992; Kosslyn et al., 1993, 1995, 1999). Consistent with the present data, a recent study reported that visual cortex activity was correlated with reported vividness, demonstrating
that individual differences in mental imagery vividness are detectable by means of fMRI (Amedi et al., 2005; Cui et al., 2007).
For tactile imagery, the right post-central gyrus (BA2) was
found activated in high-vivid subjects as compared to low-vivid
ones, as well as correlated with the vividness ratings. This lateralization is probably not due to the handedness of the subjects, since
we recruited only right-handed subjects, nor to a tactile tentative
exploration by hand, since we asked subjects to avoid any type
of movement during the scanning session, but more likely to the
subjective component of tactile sensation (Querleux et al., 1999).
In fact, even though Yoo et al. (2003) demonstrated an activation
of the left post-central gyrus (BA 1/2/3) when subjects imagined
a tactile stimulation on the back of the right hand (specically
when comparing tactile imagery with resting conditions), Querleux et al. (1999) found an activation mostly localized in the ipsilateral somato-sensory cortex during the imagination period of
tactile stimulation and a more strong activation of the contra-lateral cortex during the period of tactile perception.
Regarding gustatory imagery, the left anterior insula (BA13)
was found active in both the analyses that were carried out, as well
as the left middle frontal gyrus when high-vivid subjects were
compared to low-vivid ones. These results contrast with those of
Kikuchi et al. (2005) and with those of Olivetti Belardinelli et al.
(2004b), which revealed activations in the right insula during image generation of tastes, but they are in accordance with those of
Kobayashi et al. (2004), who found activations in the left and bilateral insulae during gustatory imagery and perception, respectively,
and in the middle and/or superior frontal gyri. According to these
authors, the left insula also processes higher gustatory information
by receiving a signal from the regions in the frontal cortex that may
mediate the top-down control of retrieving gustatory information from long-term memories.
Concerning kinaesthetic imagery, the pre-central gyrus (BA4)
and the anterior cerebellum were found activated in high-vivid
subjects only, as well as the pre-central gyrus (BA6) with a correlation analysis. Once more, for motor imagery too, the literature
provides contrasting results. Jahn et al. (2004) did not report
any activation of motor cortices during the imagery of everyday
complex movements. However, the motor cortices, although with
different patterns of activation, were found active during the
mental representation of movements with the left or the right
hand (Dechent et al., 2004; Lotze et al., 1999; Porro et al.,
1996; Roth et al., 1996) and complex everyday movements like
eating a meal, or swimming (Szameitat et al., 2007). The anterior
cerebellar activation is consistent with the view of a temporal
processing of sensory and motor information (Ivry, 1996), suggesting that the cerebellum is able to extract kinematic information relative not only to the movement itself (Kavounoudias et al.,
2008), but also to kinaesthetic imagery as well. Considering the
debate as to whether the primary motor area may be recruited
during kinaesthetic imagery, and the evidence that the Post-central Gyrus (BA 3) was found active as well as the BA 4, this activation revealed in the present study must be interpreted with
caution.

Finally, the somatic imagery modality has shown, in high-vivid

as compared with low-vivid subjects, the recruitment of the right
post-central gyrus (BA2) with a cluster size of three voxels only
and a correlation with the right post-central gyrus (BA3). Although
no other study has been carried out on this modality except those
of Olivetti Belardinelli et al. (2004a, 2004b), besides in agreement
with the results in the present study, the somato-sensory activation (post-central gyrus) yielded by the somatic imagery system
is in line with studies involving vibro-tactile stimulation (Leblanc,
Mayer, Zatorre, Tampieri, & Evans, 1995).

5.3. Modality-specic deactivations

The analysis comparing the low-vivid subjects with the high-vivid ones did not show any signicant activation in modality-specic areas. As shown by the histograms of effect sizes (Fig. 3), the
low-vivid subjects even reported deactivations in the same areas
that are vice versa activated by the high-vivid subjects.
Deactivations are normally observed in a rival cortical area
during mental imagery (Amedi et al., 2005). Nevertheless, Buckner, Raichle, Miezin, and Peterson (1996) found that occipital
deactivations can occur when people generate visual images from
verbal cues in order to recall previously learned objects. According to the authors, these deactivations reect a cross-modal inhibition process, given the auditory-directed attention paid to the
words. Interestingly, Mellet et al. (2000) found a signicant decrease of rCBF in the primary visual area when subjects were
asked to perform on a high-resolution imagery task recollecting
material visually, rather than verbally. Although in this case the
authors explicitly claim that it is hard to account for such a result,
they attempt to explain this puzzling deactivation referring to it
as a top-down modulation. In the present study, since imagery
was triggered by auditory sentences, it is possible to interpret
the deactivations in our low-vivid subjects as caused by a crossmodal inhibitory mechanism involving a verbal attention
processing.
Anyway, this pattern of results challenges the hypothesis
whether low-vivid subjects use a completely different network
with respect to the high-vivid ones. Therefore, the distinction between simulation account of conceptual knowledge processing
and mental imagery has to be taken into consideration. As
pointed out extensively by Barsalou (1999, 2008a, 2008b) and
Barsalou, Santos, Simmons, and Wilson (2008), simulation representing conceptual knowledge occurs as re-enactment of perceptual, motor, and introspective states acquired during a particular
experience. Mental imagery is a conscious simulation in working
memory, whereas, other forms of simulation are unconsciously
outside working memory (Barsalou, 1999, 2008a). Since in the
present experiment subjects were explicitly instructed to perform
a mental imagery process, the simulation account ts well with
the results obtained from our high-vivid subjects. This is not
the case for our low-vivid subjects results, which did not show
any modality-specic activation. More likely, given the deactivation patterns, the low-vivid subjects produced conscious semantic
representations based on available linguistic information, rather
than unconscious simulations partially reactivating perceptual
experiences.
Therefore, it is possible that low-vivid subjects are more condent in a conscious verbal-semantic strategy to achieve imagery
generation and use their knowledge of physical principles in the
real world to simulate how things would appear under actual
physical conditions corresponding to the tasks (Pylyshyn, 2002).
Although this interpretation may be consistent with the propositional view, further studies are necessary to better address this
issue, as well as the nature of deactivations.

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M. Olivetti Belardinelli et al. / Acta Psychologica 132 (2009) 190200

Appendix A (continued)

6. Conclusions
According to our hypothesis, the level of imagery vividness in
different sensory modalities may be related to differences of BOLD
activity in modality-specic cortices. In other words, some of the
neural processes underlying modality-specic perception may also
be used in imagery when people are able to evoke vivid images. In
particular we found modality-specic activation for visual, tactile,
gustatory, kinaesthetic and somatic imagery modalities when subjects generated and maintained mental images triggered by auditory sentences. These results are consistent with the view that
the ability of generating vivid images may have an inuence on
the process of image formation on both the format and the activation levels, high-vivid subjects creating more analogical images
and sharing very similar neural mechanisms with perception as
compared to low-vivid subjects. In the future, in order to better
specify this difference we are planning to compare the fMRI results
of high- and low-vivid subjects with their ratings, after the fMRI
session, of the perceived vividness of the image generated by each
stimuli sentence.

Appendix A
Visual
sentences

Auditory
sentences

Tactile sentences

Gustatory
sentences

To see a bulb

To hear a
whistle
To hear a
rumble
To hear a
tinkling
To hear a snap

To touch something
grainy
To touch something
velvety
To touch something
sharp
To touch something
sticky
To touch something
liquid
To touch something
hard
To touch something
smooth
To touch something
gelatinous
To touch something
soft
To touch something
rough
To touch something
gummy
To touch something
greasy

The salty taste

To see a
bucket
To see a coin
To see a
cloud
To see a book

To hear grating

To see a glass

To hear a shot

To see a box

To hear ringing

To see a hat

To hear
thunder
To hear a
chime
To hear
buzzing
To hear a
rustle
To hear an
uproar

To see a
candle
To see a
pencil
To see a
house
To see a tree

The sweet taste

The sour taste
The rancid taste
The sweet and
sour taste
The spicy taste
The acidic taste
The tart taste
The bitter taste
The fruity taste
The peppery
taste
The unripe taste

Olfactory
sentences

Kinaesthetic
sentences

Somatic sentences

Abstract
sentences

The stale
smell
The smell of
sewage
The burning
smell
The smell of
alcohol
The smell of
gas
The smell of
paint
The smell of
leather

The action
writing
The action
jumping
The action
painting
The action
walking
The action
kicking
The action
pulling
The action
lifting

The sensation
shivers
The sensation
excitement
The sensation
irritation
The sensation
pain
The sensation
fatigue
The sensation
creeps
The sensation
exertion

of

The legal
technicality
The power of
reason
Syntactical
correctness
The spiritual
power
The cardinal sin

of the

Racial prejudice

of

Innate goodness

of
of
of
of
of
of
of

of
of
of
of

Olfactory
sentences

Kinaesthetic
sentences

Somatic sentences

Abstract
sentences

The smell of
lavender
The clean smell

The action of
pushing
The action of
throwing
The action of
running
The action of
swimming
The action of
dancing

The sensation of
cramps
The sensation of cold

Formal logic

The smell of
bread
The fried smell
The moldy smell

The sensation of
nausea
The sensation of
drunkenness
The sensation of
freshness

The lack of
honesty
Universal
domination
Classical
physics
Judging on
intention

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