BRAIN IMAGING NEUROREPORT

Music in minor activates limbic structures:

a relationship with dissonance?
Anders C. Greena,c,d, Klaus B. B
rentsena,c, Hans Stdkilde-Jrgensenc, Mikkel Wallentind,
Andreas Roepstor¡ b,d and Peter Vuustd,e

a
Departments of Psychology, bAnthropology, University of Aarhus, cMR Research Centre, dCenter of Functionally Integrative Neuroscience,
Aarhus University Hospital and eRoyal Academy of Music, Aarhus, Denmark

Correspondence to Anders C.Green, Department of Psychology, University of Aarhus, Jens Chr. Skous Vej 4, DK- 8000 Aarhus C, Denmark
Tel: + 45 89424900; fax: + 45 89424901; e-mail: anders@psy.au.dk

Received 4 February 2008; accepted 13 February 2008

Using functional magnetic resonance imaging, we contrasted ma-
jor and minor mode melodies controlled for liking to study the
neural basis of musical mode perception. To examine the in£uence
of the larger dissonance in minor melodies on neural activation dif-
ferences, we further introduced a strongly dissonant stimulus, in
the form of a chromatic scale. Minor mode melodies were evalu-
ated as sadder than major melodies, and in comparison they caused
increased activity in limbic structures, namely left parahippocam-
pal gyrus, bilateral ventral anterior cingulate, and in left medial pre-
frontal cortex. Dissonance explained some, but not all, of the
heightened activity in the limbic structures when listening to minor
mode music. NeuroReport 19:711^715
c 2008 Wolters Kluwer
Health | Lippincott Williams & Wilkins.

Keywords: dissonance, emotion, functional magnetic resonance imaging, limbic, mode, music

Introduction neutral music, and pointed again to left posterior cingulate

Music has a strong effect on human emotions, which is
probably a fundamental reason for its existence in every
human culture [1]. A clear emotional dichotomy has
consistently been found at the behavioural level for Western
listeners when comparing major to minor mode music.
Major is usually referred to as the happier and minor the
sadder mode [2]. Musicologically, a main difference
between major and minor mode is that minor allows for
more dissonance than major mode. This is evident in the
array of notes utilized – and perceived as fitting by listeners
– in minor compared with major mode contexts in Western
music [3,4]. Perceived degree of dissonance potentially
influences liking and the perception of emotional content in
music [5,6]. Therefore, dissonance is a possible explanatory
factor concerning mode-related neural activation differ-
ences. This putative interplay and possible overlap between
the neural correlates of mode and of dissonance have not
been widely examined before, but is explored in this
study.
Recently, a few studies [7–10] have investigated the
impact on brain activity of major and minor mode, as well
as the related topic of happy and sad music, all using
functional magnetic resonance imaging (fMRI), but their
findings only partially overlap, and in some cases contradict
each other. One study found that listening to minor mode
excerpts, opposed to major, caused differential activity in
left posterior cingulate, and left orbitol and dorsolateral
prefrontal cortex [7]. Another study implicated the amyg-
dala, retrosplenial region, brainstem and cerebellum in a
similar contrast [8]. A third study contrasted sad with
gyrus, as well as left middle frontal gyrus, temporal
auditory areas and the hippocampus, amygdala and
cerebellum [9]. However, this study did not find any
activation when instead contrasting sad with happy music,
whereas [7] showed no activation in an opposite contrast
(major over minor). Finally, [10] indicated only extraneous
(motor related) activity when contrasting major and minor
chords, and implicated similar areas (bilateral inferior
frontal gyri, bilateral dorsal anterior cingulate and thala-
mus) when comparing both major and minor chords to
neutral ones.
Using fMRI, we contrasted major and minor mode
melodies to study the neural basis of musical mode
perception. To examine the influence of dissonance on
neural activation differences, we introduced a strongly
dissonant stimulus with minimal melodic content, in the
form of a chromatic scale. We further intended to control for
potential confounds in the earlier studies [7–10]. First,
unlike [7,9] who used preexisting music, stimuli used in our
study were purpose-made, and thus unknown to partici-
pants. This eliminated the risk that participants would
recognize some melodies but not others; an aspect known
to alter the perception and appreciation of them [11,12].
Second, in contrast to [7,9], purpose-made melodies
rendered it possible to keep them deliberately similar along
a range of parameters, such as amplitude, timbre and tempo
– all of which are known to influence music perception and
hence its neural correlates [12]. The two remaining previous
studies, in contrast, used either single chords [8], or groups
of four triads [10], as stimuli. Although highly controllable,

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 NEUROREPORT
such stimuli lack ecological validity, given that real music
unfolds over time. Third, we controlled liking for the music,
as a liking difference can by itself alter brain activation
[5,6,13].
By ensuring this high level of experimental control, and
by investigating the possible relationship of mode with
dissonance, this study aimed to clarify the neural under-
pinnings of musical emotion.
Methods
Stimuli
A pool of 30 unknown single-voice melodies of 13 s duration
each was used; half in major mode, half in minor. Melodies
were digital piano sequences, quantized and generated in
Cubase SX (Steinberg Media Technologies GmbH, Ham-
burg, Germany), postprocessed in Adobe Audition. Three
different keys (A flat, E and C) were used to add variety;
other parameters were kept identical, including ambitus
(a 10th), tempo (80 BPM) and amplitude.
The happiness or sadness expressed by the melodies was
judged by a separate group of 21 volunteers, on a scale of
1 (saddest) to 5 (happiest).
An ascending chromatic scale of uniformly distributed
dissonant intervals (minor seconds), but otherwise with
similar characteristics as the melodies, was used as the third
stimulus type.
Procedure and participants
Twenty-one healthy nonmusicians, right-handed (self re-
port), with a mean age of 27.3 (range: 20–33) were instructed
to visually fixate on a cross on a screen, listen carefully to
the stimuli over headphones, and immediately after each
melody to rate how well they liked it on a scale from 1 (least
liked) to 5 (most liked), using a response box with one
button for each finger. Stimulus sequences were pseudo-
randomized for each participant individually, and consisted
of 12 major and 12 minor mode melodies, six chromatic
scale repetitions and a number of 2–8 s silences in between.
Participants performed a practice session before the actual
fMRI scan to familiarize themselves with all aspects of
the procedure. The ethical committee of Aarhus County
approved the study.
The reported study was a separate part of a larger
investigation (forthcoming) of memory effects on music
perception. This involved a learning phase preceding the
fMRI scan, where participants listened to 18 of the melodies
a varying number of times each. In the scanner, they listened
to these again once each, along with a further six melodies
(for a total of 24). As the distribution of major and minor
mode was evenly balanced in every group of melodies,
these additional elements were considered to be of no
consequence to the results reported here.
Image acquisition and data analysis
Scanning was performed on a Signa Excite 1.5 T MR scanner
(General Electric Medical Systems, Milwaukee, Wisconsin,
USA). T1-weighted anatomical images were acquired, and
functional bold images were obtained using a gradient-
echo echoplanar imaging sequence; TR¼2700 ms, TE¼40 ms.
Thirty-four axial slices of 5-mm thickness (no gap), covering
the whole brain were acquired (64  64 matrix).
Functional MRI data analysis was done with SPM5
(Institute of Neurology, London, UK). Data preprocessing
712 Vol 19 No 7 7 May 2008
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
GREEN ETAL.
included realignment to the first image and unwarping,
coregistration with the T1-weighted images and slice
timing. Subsequently, segmentation of grey and white
matter, together with spatial normalization, was performed
using the unified segmentation approach [14], and images
were smoothed (10-mm full width at half maximum
Gaussian kernel).
Experimental effects were estimated with a general linear
model [15]. Second-level random effects analyses yielded
statistical parametric maps (SPMs). Contrasts were set up
to test for mode effect (minor vs. major). SPMs were
thresholded at Po0.05, Family Wise Error corrected for the
entire volume at voxel level, whereas cluster level correction
was also investigated. Voxel coordinates were transformed
(http://imaging.mrc-cbu.cam.ac.uk/imaging) from Montreal
Neurological Institute to Talairach space. Peak–voxel ana-
lyses were performed on the areas found in the second-level
analysis; averaged b values for the main effects of major and
minor were compared with those for the chromatic scale.
Results
Behavioural results
Minor mode melodies were judged as sadder than major:
mean major 2.62 (SD¼0.33) and mean minor 3.57 (SD¼0.40;
paired t-test: T¼8.001, Po0.001).
Major and minor mode melodies were rated as equally
likeable, mean major 3.06 (SD¼0.38) and mean minor 3.26
(SD¼0.48; paired t-test: T¼1.935, P40.05).
Functional MRI results
The contrast of minor over major melodies revealed
activation in left parahippocampal gyrus (LPHG) (Fig. 1a).
A further activation site (Fig. 1b) at cluster level correction
encompassed left and right ventral anterior cingulate cortex
(VACC) (BA 24), and extended into left medial frontal gyrus
(LMFG) of the medial prefrontal cortex (BA 10). The
opposite contrast of major over minor yielded no significant
activation.
In the peak–voxel analysis, the mean response to the
chromatic scale fell between that of major and minor mode
melodies with regard to three of the four areas mentioned
above (Fig. 2). None of these differences between chromatic
scale and the melodies were statistically significant (re-
peated measures analysis of variance, P40.05). In LMFG,
the chromatic scale elicited the greatest (least negative)
response, followed by minor, and then major mode. Here,
the difference between chromatic scale and major was
significant (Po0.05).
Discussion
In this study, we found limbic (LPHG, bilateral VACC) and
limbic-related structures (LMFG) differentially activated by
listening to melodies in minor mode, compared with
equivalent major melodies. At the behavioural level, we
confirmed that the presented melodies in minor were
perceived as sadder than those in major mode. This is
consistent with the fact that limbic structures are central to
processing emotions [16]. We further propose that degree
of dissonance may account for some but not all of the
heightened activity of the brain regions responsive to minor
mode melodies. Thus, the study conclusively shows that
NEUROIMAGING OF MUSICAL EMOTION NEUROREPORT

(a) (b)

6 6
5 5
4 4
3 3
2 2
1 1
0 0
Fig. Talairach Z score P
coordinates
(x,y,z)
(a) Voxel-level significant Voxel
activations (FWE) level
L parahippocampal gyrus −18,−44,8 4.65 0.039
(b) Cluster-level significant Cluster
activations level
R anterior cingulate (BA 24) 8, 33, 6 3.76 0.031
L anterior cingulate (BA 24) −2, 37, 6 3.70 0.031
L medial frontal gyrus (BA 10) −14, 47, 3 3.68 0.031

Fig. 1 Statistical activation maps for the minor^ major contrast of the random e¡ects analysis, withTalairach coordinates for the peak activation sites,
and their Z scores and P values.The right side of the images corresponds to the right hemisphere. FWE, Family Wise Error.

(a) 0.1 (b) 0.1 limbic structures are more involved in processing minor
than major mode melodies.

Minor versus major mode

0.0 Our finding of LPHG is unique when compared with
previous neuroimaging studies on musical mode [7–10].
0.0 However, activation of LPHG is consistent with neuroana-
tomical evidence showing functional connection between
−0.1 LPHG and the other activated regions, namely cingulate
cortex and frontal cortical areas, including BA 10 [17].
Interestingly, other neuroimaging studies have implicated
parahippocampal gyrus in listening to unpleasant music
−0.1 −0.2 [5,6], and patients with lesions of the area show decreased
Major sensitivity towards it [18]. The involvement of LPHG in this
(c) 0.1 (d) 0.1 Minor study cannot be because of unpleasantness, as the beha-
Chromatic vioural results showed no difference in liking between
minor and major. However, our results show that minor
0.0 melodies are judged as sadder than major. This indicates
0.0 that LPHG activation is not always explained by variations
in liking, but can, as in our case, also be attributed to the
−0.1 emotional valence of sadness. Furthermore, in the context of
this study it is important that the perceived unpleasantness
−0.1 of the music in the above-mentioned studies [5,6,18] was
−0.2 achieved by the use of dissonance (see below).
Activation of the ventral anterior cingulate cortex is in
general related to affective processing, whereas the dorsal
−0.3 −0.2 part is associated with cognition [19]. Furthermore, evidence
suggests that VACC is especially sensitive to negative or sad
Fig. 2 Responses to major, minor and chromatic scale (dissonance) at emotional content [20]. Involvement of VACC in this study
peak coordinates found in the minor^ major contrast. Graphs represent
averaged b values across 21participants for the three conditions in: LPHG
might thus reflect the finding that minor mode melodies
(a), right VACC (b), left VACC (c) and LMFG (d). Error bars represent were perceived as sadder than major ones. However, none
standard error. LMFG, left medial frontal gyrus; LPHG, left parahippo- of the previous imaging studies of mode effects have shown
campal gyrus; VACC, ventral anterior cingulate cortex. VACC in minor–major contrasts [7–10]. On the contrary,

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 NEUROREPORT
anterior cingulate activation was found when participants
listened to happy versus neutral music [9], and has been
shown to correlate with intensity of musical ‘chills’ [21]. It is
conceivable that the care taken in our study to optimize
experimental control, compared with earlier studies [7–10],
has allowed for a finer differentiation of anterior cingulate
activity in response to musical emotion.
The involvement of LMFG possibly reflects its strong
connectivity with the anterior cingulate cortex and other
limbic structures; a connectivity that has supported the
proposed role of the medial prefrontal cortex as a centre for
integration of emotional content from these regions [19,22].
This was corroborated with respect to musically induced
emotions by a recent study [23], which investigated the
emotionally enhancing effect of presenting congruent
classical music together with affective pictures (as opposed
to pictures alone). Both the medial frontal gyrus and the
parahippocampus were among the brain regions differen-
tially activated by the combined visual/auditive stimuli.
Additionally, activation of left medial prefrontal cortex
(BA 10) has been shown before in a minor–major contrast
[7]. The authors claimed that involvement of BA 10 was
caused by a hypothesized need to introspectively evaluate
the emotional (happy–sad) content. However, it is unclear to
us why this introspection should be more pronounced when
judging minor rather than major mode music. Conse-
quently, there is need for another explanation of what
causes the increased emotional brain activity when listening
to minor mode music in particular.
Dissonance as an explanatory factor
We tested whether the limbic activation in the minor–major
contrast might reflect the higher degree of dissonance in
minor melodies. This was performed by comparing the
response to a chromatic scale, which is strongly dissonant,
to major and minor mode melodies in the peak voxels of the
brain areas activated in the minor–major contrast. Musico-
logically, the major, minor and chromatic stimuli reflect
three different levels of musical dissonance, where major
mode is the least dissonant and the chromatic scale the
most, as all its melodic intervals are dissonant. Although
dissonance is usually associated with chords, it is also
relevant for intervals in single-voice melodies, such as our
stimuli, because subsequent notes are related to each other
in the sensory memory of the listener. Dissonance can
induce musical tension, which according to musicology is a
main carrier of musical meaning and emotion [1,24]. The
higher degree of tension in the minor mode could also be a
contributing factor to the emotional quality of sadness it
elicits in listeners, including those in our study. To test
whether dissonance alone could explain the minor–major
contrast, it was hypothesized that the chromatic scale would
elicit the strongest response, minor mode the second
strongest, and major mode the weakest in the brain areas
responding more to minor than major mode music. This
should hold true for LPHG in particular, as this area has
repeatedly been implicated in perceiving dissonance
[5,6,18], but also for the other three brain areas, in as much
as they may constitute a common functional network.
Moreover, the medial frontal gyrus and anterior cingulate
have also been involved in processing dissonance [25].
The peak–voxel analysis showed that for LPHG and the
two VACC areas, the mean response to the chromatic
714 Vol 19 No 7 7 May 2008
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GREEN ETAL.
stimulus was in between and not significantly different from
the response to the major and minor melodies (Fig. 2).
Hence, harmonic dissonance may be a contributing but
certainly not an exclusive factor in the observed minor-
related activity increase. Only in LMFG did we find traces of
the hypothesized response pattern. Musicologically, these
findings suggest that there must be more to the minor mode
than its comparatively high degree of dissonance. It is worth
emphasizing here that the dissonant stimulus material was
a simple chromatic scale, whereas the minor (and major)
mode stimuli were actual melodies. This indicates that in
order for dissonance to exert its maximum influence on
brain activity, and hence perception, it needs to occur within
a ‘real’ musical context. An obvious possibility for future
investigation would be to study dissonant melodies, not just
scales.
Conclusion
This study showed that left parahippocampal gyrus is
crucially involved when people listen to melodies in the
sad minor mode, as opposed to the happier major mode.
Further areas implicated were bilateral ventral anterior
cingulate and left medial prefrontal cortex. Compared with
earlier studies, the present design controlled liking for the
melodies, as well as a range of acoustic and musical
parameters. The greater dissonance of minor than major
may account for some, but not all, of the specific brain
activation related to listening to music in minor. The most
pertinent behavioural explanation for the observed brain
activity is the emotional quality of sadness associated with
the minor mode.
Acknowledgements
The authors thank Bo Sommerlund, Torben E. Lund, Kim
Mouridsen, Christina Scheel Meyer and Pernille Bruhn. The
study was partly funded by The Danish Research Council
for Communication and Culture, and The Danish National
Research Foundation.
References
1. Meyer LB. Emotion and meaning in music. Chicago: University of Chicago
Press; 1956.
2. Hevner K. The affective character of the major and minor mode in music.
Am J Psychol 1935; 47:103–118.
3. Krumhansl C, Kessler EJ. Tracing the dynamic changes in perceived tonal
organization in a spatial representation of musical keys. Psychol Rev 1982;
89:334–368.
4. Aarden B. Dynamic melodic expectancy. PhD dissertation. Ohio State
University; 2003.
5. Koelsch S, Fritz T, von Cramon DY, Müller K, Friederici A. Investigating
emotion with music: an fMRI study. Hum Brain Mapp 2006; 27:239–250.
6. Blood AJ, Zatorre RJ, Bermudez P, Evans AC. Emotional responses to
pleasant and unpleasant music correlate with activity in paralimbic brain
regions. Nat Neurosci 1999; 2:382–387.
7. Khalfa S, Schön D, Anton J-L, Liégeois-Chauvel C. Brain regions involved
in the recognition of happiness and sadness in music. Neuroreport 2005;
16:1981–1984.
8. Pallesen KJ, Brattico E, Bailey C, Korvenoja A, Koivisto J, Gjedde A,
et al. Emotion processing of major, minor, and dissonant chords:
a functional magnetic resonance imaging study. Ann N Y Acad Sci 2005;
1060:450–453.
9. Mitterschiffthaler MT, Fu CHY, Dalton JA, Andrew CM, Williams SCR.
A functional MRI study of happy and sad affective states induced by
classical music. Hum Brain Mapp 2007; 28:1150–1162.
10. Mizuno T, Sugishita M. Neural correlates underlying perception of
tonality-related emotional contents. Neuroreport 2007; 18:1651–1655.
NEUROIMAGING OF MUSICAL EMOTION
11. Peretz I, Gaudreau D, Bonnel A-M. Exposure effects on music preference
and recognition. Mem Cognit 1998; 26:884–902.
12. Peretz I, Zatorre RJ. Brain organization for music processing. Annu Rev
Psychol 2005; 56:89–114.
13. Menon V, Levitin DJ. The rewards of music listening: response and
physiological connectivity of the mesolimbic system. Neuroimage 2005;
28:175–184.
14. Ashburner J, Friston KJ. Unified segmentation. Neuroimage 2005; 26:
839–851.
15. Friston KJ, Holmes A, Worsley KJ, Poline JB, Frith CD, Frackowiak RSJ.
Statistical parametric maps in functional imaging: a general linear
approach. Hum Brain Mapp 1995; 2:189–210.
16. LeDoux JE. Emotion circuits in the brain. Annu Rev Neurosci 2000;
23:155–184.
17. Nolte J. The human brain: an introduction to its functional anatomy. 5th ed.
St Louis: Mosby; 2001.
18. Gosselin N, Samson S, Adolphs R, Noulhiane M, Roy M, Hasboun D,
et al. Emotional responses to unpleasant music correlates with damage to
parahippocampal cortex. Brain 2006; 129:2585–2592.
Vol 19 No 7 7 May 2008 715
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
NEUROREPORT
19. Bush G, Luu P, Posner MI. Cognitive and emotional influences in anterior
cingulate cortex. Trends Cogn Sci 2000; 4:215–222.
20. Allman JM, Hakeem A, Erwin JM, Nimchinsky E, Hof P. The anterior
cingulate cortex. The evolution of an interface between emotion and
cognition. Ann N Y Acad Sci 2001; 935:107–117.
21. Blood AJ, Zatorre RJ. Intensely pleasurable responses to music correlate
with activity in brain regions implicated in reward and emotion. Proc Natl
Acad Sci U S A 2001; 98:11818–11823.
22. Krawczyk DC. Contributions of the prefrontal cortex to the neural
basis of human decision making. Neurosci Biobehav Rev 2002; 26:
631–664.
23. Baumgartner T, Lutz K, Schmidt CF, Jäncke L. The emotional power of
music: how music enhances the feeling of affective pictures. Brain Res
2006; 1075:151–164.
24. Vuust P, Roepstorff A, Wallentin M, Mouridsen K, Østergaard L. It don’t
mean a thingy Keeping the rhythm during polyrhythmic tension,
activates language areas (BA47). Neuroimage 2006; 31:832–841.
25. Foss AH, Altschuler EL, James KH. Neural correlates of the Pythagorean
ratio rules. Neuroreport 2007; 18:1521–1525.