[Link].

com/scientificreports

OPEN Emotional Intent Modulates The

Neural Substrates Of Creativity: An
fMRI Study of Emotionally Targeted
received: 22 July 2015
accepted: 18 November 2015 Improvisation in Jazz Musicians
Published: 04 January 2016
Malinda J. McPherson1, Frederick S. Barrett2, Monica Lopez-Gonzalez3, Patpong Jiradejvong1
& Charles J. Limb1,3,4

Emotion is a primary motivator for creative behaviors, yet the interaction between the neural systems
involved in creativity and those involved in emotion has not been studied. In the current study, we
addressed this gap by using fMRI to examine piano improvisation in response to emotional cues. We
showed twelve professional jazz pianists photographs of an actress representing a positive, negative
or ambiguous emotion. Using a non-ferromagnetic thirty-five key keyboard, the pianists improvised
music that they felt represented the emotion expressed in the photographs. Here we show that activity
in prefrontal and other brain networks involved in creativity is highly modulated by emotional context.
Furthermore, emotional intent directly modulated functional connectivity of limbic and paralimbic
areas such as the amygdala and insula. These findings suggest that emotion and creativity are tightly
linked, and that the neural mechanisms underlying creativity may depend on emotional state.

The ability of art to serve as a means of emotional self-expression and emotional communication has arguably been
one of the fundamental reasons for the omnipresence of art throughout all cultures in human history1,2. Creative
mediums such as painting, poetry, dance, film and music evoke intense emotions for both artists and audiences
alike, allowing humans to experience and share a wide range of emotional responses within a secure framework3.
Emotion often serves as a catalyst for creative expression4,5, and therefore it is crucial to understand how emotion
impacts the neural mechanisms that give rise to creativity, and also to understand how creative artistic expression
can modulate the neural systems responsible for processing emotions. Here we examined the effects of emotional
intent on the neural systems involved in the creation of emotional music compositions using functional magnetic
resonance imaging (fMRI). We instructed professional jazz pianists to improvise short pieces in response to vis-
ually presented emotional cues with positive, ambiguous, and negative valences (Fig. 1b). Using this paradigm,
we examined whether emotional intent influences the neural substrates of creativity.
To our knowledge, this experiment represents the first effort in neuroscience to address the relationship between
musical creativity and emotional expression. The majority of the literature on both emotion and music (and emo-
tion in other domains) has examined the perception of emotions rather than the expression of emotions. This is
a major deficit in the literature, as a comprehensive model of human emotion processing must include both per-
ception and expression (production) of emotional states, two distinct yet intrinsically linked human experiences.
Since the creative expression of emotions through music and other artistic mediums is often used to regulate,
project, and alter emotional states6, art may be arguably the best means by which to first approach the complex
relationships between creativity and emotion, and how they interact neurologically.
Creativity can be broadly defined as the ability to produce something that is both novel and valuable for a
given context7. Part of the difficulty in studying and modeling creativity is that the nature of creative pursuits in
the arts is not bound by notions of correct or incorrect, right or wrong, or even better or worse; indeed violations

1
Department of Otolaryngology-Head and Neck Surgery, Johns Hopkins School of Medicine, Baltimore, Maryland,
21205, USA. 2Department of Psychiatry and Behavioral Sciences, Johns Hopkins School of Medicine, Baltimore,
Maryland, 21205, USA. 3Peabody Conservatory of The Johns Hopkins University, Baltimore, Maryland, 21202, USA.
4
Department of Otolaryngology-Head and Neck Surgery, University of California San Francisco, San Francisco,
California, 94143, USA. Correspondence and requests for materials should be addressed to M.J.M. (email:
malindajmcpherson@[Link])

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Figure 1.  fMRI Paradigm and Photographs. Used as Visual Stimuli (a) Schematic showing fMRI stimulus
and instruction presentation paradigm and (b) Photographs representing positive, ambiguous and negative
emotions. Photographs were shot indoors in black and white with a 50 mm lens at f16 using a Nikon D700
digital SLR camera.

of expectation are a key criteria used to determine whether something is considered creative8 and art critics
throughout history have harshly criticized works of art that eventually came to be regarded as masterpieces9. In
music, violations of expectation (surprises) are often a critical feature of its emotional content10–12. Improvisation
is a form of immediate creativity in which artists can spontaneously generate novel material, and is a defining
feature of jazz music. Jazz musicians are highly trained experts in improvisation, and they are able to reliably
perform emotionally targeted improvisations in a laboratory setting13. Jazz is a diverse and unrestrictive form of
music, and jazz improvisation enables musicians to flexibly incorporate many musical features in order to express
an emotion14,15. Therefore, musical improvisation offers a unique opportunity to systematically examine the effects
of emotion on creativity (and creativity on emotion) in an ecologically valid setting.
A growing number of researchers have studied neural activation during creative tasks in both musical and
nonmusical domains, however none have directly addressed the role of expressing emotion through naturalistic
improvisation16–23. Multiple studies have reported improvisation-related activations in the premotor cortex, sup-
plementary motor area (SMA), perisylvian language areas, and the medial prefrontal cortex. Some of these studies
have reported deactivations during improvisation in the dorsolateral prefrontal cortex (DLPFC) and parietal
areas, including the angular gyrus (AG) and precuneus. Deactivation in the DLPFC may indicate that creativity
can induce a state of total immersion often referred to as a “flow state”24. More widespread DLPFC deactivation,
referred to as hypofrontality, may be indicative of deeper flow states25. Creativity is not a single unified set of mental
processes or abilities. While some types of creativity may require intense concentration and thought, other forms
of creativity, such as jazz improvisation, may be predicated on “letting go”8. Hypofrontality may represent a neural
signature of these types of creative experiences25. A recent study by Pinho and colleagues23 used an emotional
constraint (happy or fearful) vs. a tonal constraint (six tonal or atonal pitches), and confirmed findings that DLPFC
activity is lower during free improvisation when compared to tightly constrained improvisation. While emotion
was a factor in their study, their goal was to examine the differences between a more loosely constrained, emotional
improvisation task, and a more constrained improvisation task, where the pitches were pre-determined. This differs
from the current study’s goal of examining differences between free improvisations with different emotional targets.
Similarities between previous studies of creativity suggest that there is a consistent functional network of
brain areas responsible for artistic creativity. Yet the existence of this functional network does not reveal how
it can produce such a diverse range of creative behaviors that are fundamentally unpredictable. Consequently,
identification of the factors that modulate this network is critically important in understanding the capacity of the
brain to generate the wide range of ideas that are characteristic of human creativity. In light of the central role of
emotion in creative pursuits, we sought to identify whether or not emotional intent modulates the neural systems
involved in creativity, possibly lending support to a dynamic neural model of creativity that is malleable in response
to numerous external influences.

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A. Minor B. Note Minima and Maxima

Mode 85
Major
*
100% *
90% 20.83% 80
80%
70% 54.17% 75
68.75%

Note Number
60%
50% 70
40% 79.17%
30% 65
20% 45.83%
31.25% 60
10%
0%
Negative Ambiguous Positive 55
D.
Note Density 50
5 *
*=p<.001
Negative
4.5 Ambiguous 45 *p<.05, for Note Maximums (above) and Note Minimums (below)

4 Positive Negative Ambiguous Positive

Notes per Second

*
3.5 C. Duration Distribution
3 * Negative Ambiguous Positive

Proportion (%)

Proportion (%)

Proportion (%)
0.4 0.4 0.4
2.5
2 0.3 0.3 0.3

1.5 0.2 0.2 0.2

1
0.1 0.1 0.1
0.5
0 0 0
0 123456789 123456789 123456789
Duration Duration Duration

Figure 2.  MIDI Analysis Results. Differences in proportion of major to minor keys, note minima and maxima,
distributions of note durations, note density.

We hypothesized that patterns of activation of the distributed network of brain regions involved in musical
improvisation (compared to a chromatic scale performance control condition) as well as functional connectivity
between this network and brain areas that support emotion processing would be modulated by emotional intent.
As flow states are generally associated with pleasurable experiences24, and performing both happy and sad music
can be intensely pleasurable26,27, we predicted that there would be stronger engagement of reward and flow state
areas of the brain during both positive and negative improvisation when compared to ambiguous improvisation.
We addressed these hypotheses using an ecologically valid model of spontaneous musical performance in expert
jazz pianists improvising in response to emotional targets during fMRI brain scanning.

Results
Music Performance Analysis.  Analysis of musical mode showed that 31.25% of the negative improvisa-
tions were in a major key compared with 79.17% of the positive improvisations and 45.83% of ambiguous improv-
isations (Fig. 2a).
A main effect of emotion was found for note minima [F(1, 3) =  4.04, p <  0.05] and note maxima [F(1, 3) =  8.49,
p <  0.001]. This was primarily due to differences between negative and positive improvisations (significant for
both maxima and minima (p <  0.001), as ambiguous improvisations were not statistically different from positive
or negative improvisation trials (Fig. 2b).
We observed statistical differences (p <  0.05) between corresponding bins of the duration distributions for
negative, positive and ambiguous improvisations in all but Bin 3 (¼ s; no significant difference between ambiguous
and positive, p < 0.05), Bin 4 and Bin 6 (approximately 7/20 s and 14/20 s respectively, with no significant differences
between ambiguous and negative, p < 0.05) (Fig. 2c).
A main effect of emotion was found for note densities between improvisation conditions [F(1, 3) =  31.88,
p <  0.001]. There was a significant difference between the note densities of positive and negative improvisation
conditions and ambiguous and positive improvisation conditions (p <  0.001). Higher note densities were used to
express positive emotions, and lower note densities were used to express negative and ambiguous emotions (mean
note densities: negative =  2.09, ambiguous =  2.56, positive =  3.61). There was no main effect of emotion for note
density during chromatic scale trials [F(1, 3) =  0.36, p =  0.68] (Fig. 2d).

Functional Neuroimaging: Contrast analyses.  Emotional intent modulates activity in brain regions
associated with improvisation. To examine brain areas involved in improvisation within each emotion condition
we completed within-emotion contrasts (ex. [PosImprov >  PosChrom]). We observed activations in left Broca’s
area (BA 45) in all emotion conditions and SMA (BA 6) for ambiguous and negative improvisation, as well as
deactivation in bilateral AG (BA 40), middle precuneus (BA 7), medial and lateral frontopolar prefrontal cortex

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Figure 3.  Within-Emotion Contrasts. Three-dimensional surface projections of activations (ex.

PosImprov > PosChrom) and deactivations (ex. PosChrom > PosImprov) during improvisation for different
emotion conditions. Results are from a random effects model, p <  0.005 with a 10 voxel cluster threshold.
Improvisation was associated with perisylvian language area activations and supplementary motor area
activations across emotions, as well as deactivations in the DLPFC, angular gyrus, and precuneus. The scale bar
shows the range of t-scores; the axes demonstrate anatomic orientation. Abbreviations: A, anterior; P, posterior;
R, right; L, left.

(BA 10), and DLPFC (BA 9) for all emotion conditions. There were deactivations in the left hippocampus during
positive contrasts, right hippocampus during ambiguous, and bilateral hippocampus during negative contrasts.
Additionally, we saw bilateral middle cingulate (BA 31) deactivations for all emotions. Though these activations
and deactivations were present among all emotions (SMA notwithstanding), the extent of the voxel clusters varied
widely between each emotion. Most notably, positive improvisation showed much more widespread deactivation
in the DLPFC, AG, and precuneus than negative and ambiguous improvisation (Fig. 3, Table 1).
These within-emotion contrasts also revealed clusters that were emotion-specific (i.e. not shared between
emotions). The [AmbImprov >  AmbChrom] contrast showed activations in the right superior temporal lobe
(BA 22 and BA 21, Wernicke’s Area homologue) and temporal pole (BA 38), but these activations were not seen
in [PosImprov >  PosChrom] or [NegImprov >  NegChrom]. Recent evidence indicates that the right hemisphere
language areas are involved in resolving lexical ambiguity28, therefore these results suggest that Wernicke’s area
right hemisphere homolog may be involved in processing musical ambiguity as well.
To further examine the effect of emotion on improvisation, we looked at the between-emotion contrasts (ex.
[PosImprov >  NegImprov]). These contrasts revealed significant differences between emotion blocks, most notably
in areas known for their involvement in emotional processing. The most extensive differences were seen when
negative and ambiguous improvisation were compared to positive improvisation. [AmbImprov >  PosImprov] and
[NegImprov >  PosImprov] contrasts both revealed heightened activity in the right insula (BA 13 and 47) and right
anterior cingulate cortex (BA 32), as well as activation in the right parietal cortex (BA 40), bilateral middle temporal
lobes (BA 22), and bilateral middle frontopolar prefrontal cortex (FPPFC, BA 10) (Fig. 4, Table 2). Positive improv-
isation showed increased cerebellar activity compared to negative improvisation ([PosImprov >  NegImprov]),
and markedly increased activity in the left hippocampus and amygdala and right parahippocampal gyrus when
compared to ambiguous improvisation ([PosImprov >  AmbImprov]) (Fig. 4, Table 2).
In addition to the differences described above, we observed differences between negative and ambiguous
improvisation trials. The contrast [NegImprov >  AmbImprov], showed increased activity in the right anterior
cingulate (BA 9), left angular gyrus and supramarginal gyri (BA 39 and 40) and the right hippocampus. The
converse contrast ([AmbImprov >  NegImprov]) showed a relative increase in activity in the right cerebellum
(Vermis, VII), bilateral Heschl’s gyrus (BA 41), and left primary motor areas (BA 4) during ambiguous improv-
isation. The observed activity in the cerebellum and primary motor area may be due to the increased number of
notes played (increased movement) during ambiguous improvisation compared to negative improvisation. For
between-emotion viewing contrasts, there were no significant clusters between any of the emotion and viewing
conditions, [AmbView >  PosView], [PosView >  AmbView], [PosView >  SadView], [PosView >  NegView], or
[NegView >  AmbView]. There were small ( <  16voxel) significant clusters for the contrast, [AmbView >  NegView],
in the insula, caudate and cerebellum (Cerebellum—6). The minimal number of significant differences between
viewing conditions suggests that widespread differences between emotion improvisation conditions cannot be
attributed to the emotional valence of the cue photographs. The photographs, rather than eliciting strong emotions,
were serving as emotional ‘guideposts’ for the subjects.

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Activations Negative Improvisation

BA Left Hemisphere Right Hemisphere
Region t-score x y z Cluster Size t-score x y z Cluster Size
SMA 6 5.47 − 4 0 66 145* 4.61 4 14 64 145*
Precentral Gyrus 9 6.1 − 52 12 32 42 — — — — —
Perisylvian Language Areas
IFG-P-OP 44 5.07 − 54 10 16 48 — — — — —
IFG-P-Tr 45 4.57 − 54 10 20 11 — — — — —
Deactivations Negative Improvisation
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Hippocampus − 3.32 − 30 − 36 − 8 19 − 3.29 32 − 16 − 14 13
Prefrontal Cortex
FPPFC 10 − 4.77 − 34 58 2 35 − 3.88 40 50 4 30
DLPFC 9 − 3.39 − 30 30 38 14 — — — — —
Secondary Clusters 9 − 4 − 44 18 40 13 — — — — —
DMPFC 8 − 3.93 − 6 36 48 26 — — — — —
9 − 3.46 − 4 50 24 10 — — — — —
Cingulate Gyrus
MCC 31 − 5.29 − 2 − 42 36 197* − 4.7 4 − 42 32 197*
Parietal Lobe
Superior Parietal 40 − 3.97 − 54 − 58 38 129 − 4.09 48 − 64 46 40
AG 39 − 4.12 − 52 − 60 24 83 − 3.95 48 − 66 42 11
Precuneus 7 − 4.12 − 8 − 64 38 86* − 3.72 6 − 64 38 86*
Activations Ambiguous Improvisation
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
SMA 6 4.59 − 2 12 60 24 — — — — —
Secondary Clusters 6 3.85 − 2 − 8 62 37 — — — — —
Perisylvian Language Areas
IFG-P-OP 44 4.67 − 54 8 12 64 3.96 52 16 12 21
IFG-P-Tr 45 6.63 − 50 18 16 19 4.12 54 18 15 33
STG 22 — — — — 5.89 54 − 26 0 30
Deactivations Ambiguous Improvisation
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Hippocampus − 5.35 28 − 16 − 20 52
Prefrontal Cortex
FPPFC 10 − 5.14 − 28 54 18 102* − 5.13 10 64 14 102*
DLPFC 9 − 3.87 − 36 22 40 49 − 3.98 46 24 40 10
Secondary Clusters 9 − 3.87 − 38 34 42 24 — — — — —
DMPFC 8 — — — — — − 4.42 24 22 42 38
9 − 4.08 − 14 40 20 15 — — — — —
Cingulate Gyrus
MCC 31 − 3.84 2 − 38 36 70* − 5.07 − 2 − 38 36 70*
ACC 32 − 5.69 8 38 28 62
Parietal Lobe
Superior Parietal 40 − 7.81 − 54 − 56 42 136 − 4.9 50 − 54 48 11
AG 39 − 5.59 − 52 − 66 32 74 − 4.97 46 − 70 34 21
Precuneus 7 − 3.56 − 4 − 60 42 160* − 4.66 8 − 62 44 160*
Activations Positive Improvisation
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Precentral Gyrus 9 5.24 − 56 10 34 78 — — — — —
Perisylvian Language Areas
IFG-P-OP 44 3.16 − 52 10 22 20 — — — — —
Deactivations Positive Improvisation
Region BA Left Hemisphere Right Hemisphere

Continued

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Activations Negative Improvisation

BA Left Hemisphere Right Hemisphere
Region t-score x y z Cluster Size t-score x y z Cluster Size
Hippocampus − 9.8233 − 26 − 36 − 2 236
Prefrontal Cortex
FPPFC 10 − 4.01 − 25 60 2 − 4.85 32 56 2 52
DLPFC 9 − 3.16 − 30 32 42 13 − 5.62 38 24 40 234
Secondary Clusters 9 — — — — — − 7.24 32 26 46 190
DMPFC 8 − 4.31 − 2 30 54 107* − 3.93 2 30 52 107*
9 − 3.81 − 6 46 20 166* − 4.31 6 46 38 166*
Cingulate Gyrus
MCC 31 − 5.32 − 2 − 44 34 470* − 5.32 4 − 38 34 470*
ACC 32 − 3.89 − 8 42 16 71* − 3.81 6 42 16 71*
Parietal Lobe
Superior Parietal 40 − 6.86 − 56 − 58 30 258 − 7.43 60 − 48 40 627
AG 39 − 7.01 − 58 − 88 28 229 − 3.51 58 − 58 24 64
Precuneus 7 − 5.82 − 2 − 64 36 569* − 7.47 10 − 64 32 569*

Table 1.  Within-Emotion Contrasts. All coordinates are described according to the Montreal Neurological
Institute system, and were obtained through a random effects (uncorrected, p < 0.005, 10 voxel cluster
threshold) analysis of data. Abbreviations: SMA, supplementary motor area; IFG-P-Op, inferior frontal gyrus
pars opercularis; IFG-P-Tr, inferior frontal gyrus pars triangularis; STG, superior temporal gyrus; FPPFC,
frontopolar prefrontal cortex; DLPFC, dorsolateral prefrontal cortex; DMPFC, dorsomedial prefrontal cortex;
MCC, middle cingulate cortex; ACC, anterior cingulate cortex; AG, angular gyrus.

Functional Connectivity: Psycho-Physiological Interaction (PPI) Analysis.  We originally hypoth-

esized that functional connectivity between brain areas that support emotion processing and regions of the brain
that support improvisation would be modulated by emotional intent. Thus, we looked for regions of the brain that
support emotion processing to use as seeds in a PPI analysis. Due to the prominent role of the amygdala and ante-
rior insula in previous investigations of music-evoked emotions29 and more general emotional experience30, the
left amygdala and left anterior insula clusters from the between-emotion contrasts were selected as seed regions
for PPI analysis. Prefrontal regions, such as the DLPFC, were not chosen as seeds related to emotion processing,
as these regions, even when they appear in studies of emotion processing, are typically interpreted as supporting
cognitive processes that are secondary to emotional experience or responsive to task demands, rather than pri-
mary to a given emotional experience30. Within-emotion contrasts in the amygdala PPI analysis demonstrated
lower connectivity during positive improvisation than during the positive chromatic condition between the left
amygdala and the cerebellum (Table 3a). A decrease was observed during negative improvisation compared to the
negative chromatic condition in amygdala connectivity with the right inferior frontal gyrus and left postcentral
gyrus areas (Table 3a, Fig. 5a).
In the between-emotion contrasts, overall greater connectivity in the positive improvisation condition than in
the negative improvisation condition was observed between left amygdala and the superior medial and superior
frontal gyri, the anterior cingulate, the supramarginal gyrus, and the inferior parietal sulcus (Table 3a, Fig. 5a).
Lower effective connectivity in the positive improvisation condition than in the positive chromatic condition was
observed between the left insula and superior and middle frontal gyri, precentral and postcentral gyri, and the
supramarginal gyrus, and greater effective connectivity was observed between the left insula and primary visual
cortex. We also observed lower effective connectivity between the left insula and middle frontal gyrus, and the
left insula and inferior parietal lobule, with greater effective connectivity between left insula and superior medial
gyrus. When considering the between-emotion contrast, greater effective connectivity was observed between the
left insula and the rolandic operculum during positive improvisation than during negative improvisation. Greater
effective connectivity was also observed between the left insula and a midbrain region that includes the substantia
nigra during negative improvisation compared to positive improvisation (Table 3b, Fig. 5b).

Discussion
Here we examined the interaction between emotional expression and creativity in an ecologically valid context.
MIDI performance analysis revealed that pianists were consistently using musical features such as mode, note
density, duration and range to distinguish positive, ambiguous and negative emotions. These results match findings
from a previous study of jazz improvisation in response to visual emotional cues, done outside of a scanner13, and
this similarity indicates that the pianists were improvising comfortably within the fMRI scanner. Furthermore,
the pianists were playing on a keyboard with full size keys, and they could comfortably hear their playing in real
time. It could be argued that by playing (and simultaneously hearing) their emotionally expressive improvisa-
tions, the subjects were inducing emotions in themselves through a positive auditory feedback loop. While this
is possible, a feedback effect may occur whether pianists are able to hear themselves or not, as there is evidence
for strong auditory-motor coupling in professional pianists. Several studies have shown similar neural activations
between pianists performing on a mute vs. sounding piano31,32. If hearing their improvisations increased the
musicians’ emotional arousal, this increase is in line with the aims of the current study. When we examined the

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Figure 4.  Between-Emotion Contrasts. Top row shows an axial slice rendering of contrasts between positive
and ambiguous improvisation (PosImprov > AmbImprov—Yellow) and positive and negative improvisation
(PosImprov > NegImprov—Red). Specific to positive improvisation, we observed increased activation in
the left amygdala and right cerebellum. The middle row shows sagittal slice renderings of contrasts between
ambiguous and positive improvisation (AmbImprov >  PosImprov—Cyan) and negative and positive
improvisation (NegImprov >  PosImprov—Green). These contrasts showed widespread differences, primarily
in right hemisphere superior temporal lobe, right insula, anterior cingulate cortex and superior parietal lobe.
The third row shows sagittal Axial slice renderings of contrasts between negative and ambiguous improvisation
(AmbImprov >  NegImprov—Violet, NegImprov >  AmbImprov—Blue). Primary differences between
ambiguous and negative Improvisation were increased anterior cingulate activation during negative, and
increased right superior temporal lobe and primary motor area activations during ambiguous. Sagittal sections
show axial slice location. Coronal sections show sagittal slice locations. All results are from random effects
models, p <  0.005, 10 voxel cluster threshold. Bar graphs indicate percent signal change at cluster maxima,
plotted using rfxplot, Glascher 2009). White =  positive improvisation, Dashed =  ambiguous improvisation,
Grey =  negative improvisation.

functional neuroimaging results, we found that the creative expression of emotions through music may engage
emotion-processing areas of the brain in ways that differ from the perception of emotion in music. We also
observed a functional network involved in creative performance, and the extent of activation and deactivation in
this network was directly modulated by emotional intent. Our viewing controls showed that there were few sig-
nificant differences between neural activity in response to any of the visual cues, therefore the differences between
improvisation conditions are the result of the creative expression of emotion through music, rather than a direct
response to the visual stimuli. These results highlight that creativity is context-dependent, and emotional context
critically impacts the neural substrates of artistic creativity.
Our study suggests that expressing emotion through music engages limbic and prefrontal areas of the brain in a
distinctive manner not previously observed in other studies of music perception. Although insular and hippocam-
pal activity has been observed in response to joyful music29,33–35, we observed an opposite pattern of more robust
insular and hippocampal activity during negative improvisation. A possible explanation for this difference is that
music perception generally activates the network involved in creativity (the precuneus, MPFC, DLPFC, etc.)36,37.
The deactivation of this system during creative tasks may change how the neural networks involved in emotion are

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Activations NegativeImprov > PositiveImprov
BA Left Hemisphere Right Hemisphere
Region t-score x y z Cluster Size t-score x y z Cluster Size
SMA 6 — — — — — 5.28 6 − 12 72 277
ACC 32 — — — — — 4.71 16 34 26
Insula 13 3.8 − 40 8 − 4 17 3.41 36 12 − 8 13
47 — — — — — 3.83 40 24 2 25
Frontal Lobe
FPPFC 10 3.7 − 40 40 14 10 4.11 26 46 30
9 4.19 − 42 12 38 14 4.55 38 20 34 9
Perisylvian Language Areas
MTG 21 3.98 − 64 − 40 0 22 5.36 60 − 42 − 2 62
STG 22 4.24 − 64 − 38 2 19 4.48 56 − 44 2 54
Parietal Lobe
Precuneus 7 — — — — — 5.41 4 − 44 46 136
Angular Gyrus 7 — — — — — 3.21 38 − 58 50 19
Superior Parietal 40 4.29 − 64 − 32 32 122 5.64 44 − 46 44 264
Activations AmbiguousImprov >  PositiveImprov
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
SMA 6 4.33 − 2 2 60 225* 4.59 20 − 2 64 225*
Insula 13 — — — — — 4.4 36 22 2 62
Insula 47 — — — — — 4.63 38 22 0 31
Cingulate Lobe
ACC 32 — — — — — 4.36 16 34 36 30
MCC 24 — — — — — 6.16 8 4 38 85
Frontal Lobe
Precentral gyrus 9 — — — — — 7.2 54 12 32 235
FPPFC 10 4.4 − 26 54 4 27 4.7 34 44 28 60
Perisylvian Language Areas
STG 22 4.14 − 64 − 42 2 75 6.04 66 − 14 4 124
IFG-P-OP 44 7.67 − 44 16 8 17 6.6 52 12 8 125
IFG-P-Tr 45 5.52 − 46 18 6 29 5.56 48 22 10 121
Parietal Lobe
Angular Gyrus 40 — — — — — 4.59 36 − 56 48 337
Precuneus 7 — — — — — 4.63 20 − 74 44 170
Activations PositiveImprov >  NegativeImprov
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Cerebellum—VI 4.04 − 8 − 64 − 26 113 4.07 10 − 64 − 24 157
Parahippocampal gyrus 35 — — — — — 3.51 22 − 24 − 22 7
Activations PositiveImprov >  AmbiguousImprov
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Amygdala 6.18 − 22 − 2 − 22 59 — — — — —
Hipocampus 3.9 − 32 − 26 − 14 95 — — — — —
Parahippocampal gyrus 28 — — — — — 3.32 22 − 14 − 24 11
Activations AmbiguousImprov >  NegativeImprov
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Vermis—VII — — — — — 3.94 0 − 72 − 26 17
Primary Motor Area 4 10.05 − 46 − 14 54 238 — — — — —
STG 22 4.69 − 50 − 14 4 45 5.87 56 − 8 − 2 85
Heschel's Gyrus 41 3.63 − 48 − 20 10 65 4.06 48 − 20 10 47
Activations NegativeImprov >  AmbiguousImprov
Region BA Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Hippocampus — — — — — 4.8 30 − 40 − 2 12

Continued

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Activations NegativeImprov > PositiveImprov
BA Left Hemisphere Right Hemisphere
Region t-score x y z Cluster Size t-score x y z Cluster Size
ACC 9 — — — — — 5.74 14 40 24 19
Angular Gyrus 39 3.69 − 48 − 72 30 19 — — — — —
SMG 40 3.76 − 60 − 56 28 12 — — — — —

Table 2.  Between-Emotion Contrasts. All coordinates are described according to the Montreal Neurological
Institute system, and were obtained through a random effects (uncorrected, p < 0.005, 10 voxel cluster
threshold) analysis of data. Abbreviations: SMA, supplementary motor area; IFG-P-Op, inferior frontal gyrus
pars opercularis; IFG-P-Tr, inferior frontal gyrus pars triangularis; STG, superior temporal gyrus; FPPFC,
frontopolar prefrontal cortex; DLPFC, dorsolateral prefrontal cortex; DMPFC, dorsomedial prefrontal cortex;
MCC, middle cingulate cortex; ACC, anterior cingulate cortex; AG, angular gyrus.

recruited during performance when compared to perception. These differences are critical, as they may relate in
part to the fundamentally different processes of observing an emotional state in somebody or something else and
personally experiencing that emotional state. Clearly, these processes are linked to one another through empathy
and experience, yet it is possible that the personal experience of an emotion and behavioral changes caused by the
need or desire to express that emotion (through art or otherwise) are more compelling than observation of that
emotion alone. Future studies of the key differences between emotional expression and perception are needed so
that we may arrive at a more comprehensive neural model of emotion.
Our random effects analyses indicate that there is an essential functional network involved in creative expres-
sion, and that this network is modulated by emotional intent. Several studies have shown that spontaneous musi-
cal improvisation induces deactivations in the DLPFC, angular gyrus, and precuneus, and activations in the
SMA and perisylvian language areas20–22,38–40. We observed changes in all of these areas in response to the three
different emotional intents. Most strikingly, DLPFC deactivation was much more pronounced during positive
improvisation when compared to negative and ambiguous improvisation. A decrease in prefrontal cortical activity
(hypofrontality) has been implicated in various altered states of consciousness and flow states24,25. Furthermore,
while we did not observe activations in the SMA during positive improvisation, it was active during negative and
ambiguous improvisation, primarily in the left hemisphere (expected due to the pianists’ use of their right hand
while playing). Based on the MIDI data analysis, it is unlikely that this SMA activity simply relates to increased
motor activity – performers were, on average, playing fewer notes during negative and ambiguous improvisations
when compared to positive improvisation. The SMA is often active during tasks involving continuous monitoring
of the appropriateness of motor output, including rhythmic tapping and complex sequential motor tasks41. The
observed hypofrontality and lack of SMA activity may indicate that positive improvisation induces a deeper state
of flow than negative and ambiguous improvisation.
Since listening to sad music is often pleasurable42, we originally predicted that we would observe increased
hypofrontality during negative improvisation as well as positive improvisation (when both were compared to
ambiguous improvisation). The combination of our random effects and functional connectivity analyses present
an intriguing alternative: positive and negative improvisation may be pleasurable for different reasons. Negative
improvisation showed increased functional connectivity between the insula and the substantia nigra (a dopamin-
ergic reward area) when compared to positive improvisation. This type of connectivity could imply an increased
binding of visceral awareness43, and this finding is consistent with a recent report outlining the rewarding properties
of sad music42. Music-related rewards include the ability to experience or express emotions without any “real-life”
costs. This is a reward that could certainly be maximized during music improvisation, and that could be expected
to drive activity in dopaminergic regions such as the substantia nigra. The rewarding aspects of sad music may also
be contingent on maintaining a degree of cognitive distance from the artwork3. This interpretation is consistent
with increased activation in areas responsible for cognitive control and self-monitoring, primarily the SMA and
frontal polar prefrontal cortex, during negative improvisation when compared to positive improvisation. While
further work is needed to clarify this complex relationship, it is plausible that the pleasure or satisfaction derived
from happy and sad music may be mediated through different neurobiological systems. While positive emotional
targets enable more widespread hypofrontality and deeper flow states during spontaneous creativity, negative
emotional targets may be more closely linked to a stronger visceral experience and greater activity in reward
processing areas of the brain during improvisation.
In summary, this study shows that the impulse to create emotionally expressive music may have a basic neural
origin: emotion modulates the neural systems involved in creativity, allowing musicians to engage limbic centers
of their brain and enter flow states. The human urge to express emotions through art may derive from these wide-
spread changes in limbic, reward, and prefrontal areas during emotional expression. Within jazz improvisation,
certain emotional states may open musicians to deeper flow states or more robust stimulation of reward centers.
The creative expression of emotion through music may involve more complex mechanisms by which the brain
processes emotions, in comparison to perception of emotion alone. Additional studies of how emotional state
modulates creativity in non-artistic domains such as decision-making and social interactions are needed. Future
studies could also examine whether there is an effect of gender on emotional expression through music, and
whether the neural results are altered if subjects use both hands during improvisation. This study examines just
one of many possible factors that could influence the neural underpinnings of human creativity, and there is huge
scope for investigation. Further understanding how emotion influences creativity in both artistic and non-artistic
contexts will be crucial for the derivation of a more comprehensive and accurate neural model of human creativity.

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A) Amygdala Seed (−22, −2, −22)

Region PositiveImprov >  PositiveChromatic
Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Cerebellum − 5.3549 − 8 − 38 − 22 423 — — — — —
NegativeImprov >  NegativeChromatic
Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Inferior Frontal Gyrus (pars
— — — — — − 5.7533 42 46 − 4 27
orbitalis)
Postcentral Gyrus − 6.6215 − 56 − 12 44 39 — — — — —
PositiveImprov >  NegativeImprov
Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Superior Medial Gyrus 7.9511 − 10 30 32 212 — — — — —
Superior Frontal Gyrus 7.0567 − 18 4 68 65 — — — — —
Anterior Cingulate Cortex — — — — — 7.3624 8 30 20 25
Supramarginal Gyrus — — — — — 4.9477 60 − 28 28 23
Inferior Parietal Sulcus — — — — — 5.7853 42 − 42 34 88
B) Insula Seed (− 43, 14, − 4)
PositiveImprov >  PositiveChromatic
Region Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Superior Frontal Gyrus − 5.7837 − 16 26 52 24 — — — — —
Middle Frontal Gyrus − 6.7562 − 42 22 32 221 — — — — —
*Middle Frontal Gyrus — — — — — − 6.8154 20 0 36 32
*Frontal Eye Fields — — — — — − 5.3609 30 2 36 21
Precentral Gyrus − 6.1561 − 42 − 4 30 89 — — — — —
Postcentral Gyrus − 6.5393 − 36 − 34 44 285 — — — — —
Supramarginal Gyrus − 5.0949 − 58 − 46 32 20 — — — — —
Middle Occipital Gyrus — — — — — 5.5648 26 − 88 14 55
*White matter — — — — — − 5.0431 24 44 12 25
NegativeImprov >  NegativeChromatic
Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Superior Medial Gyrus — — — — — 6.0208 6 60 18 30
Middle Frontal Gyrus — — — — — − 8.0061 32 26 22 90
Inferior Parietal Lobule − 4.8395 − 46 − 32 36 72 — — — — —
*Callosum − 8.8971 − 20 − 20 44 1178 — — — — —
PositiveImprov >  NegativeImprov
Left Hemisphere Right Hemisphere
t-score x y z Cluster Size t-score x y z Cluster Size
Rolandic Operculum — — — — — 5.1469 50 − 8 12 53
*Substantia Nigra — — — — — − 7.6075 6 − 12 − 14 29
* Cluster was not specified in the Anatomy Toolbox, and was identified using NeuroSynth. p <  0.01, k >   =  20

Table 3.  PPI Results. All coordinates are presented in the Montreal Neurological Institute template space.
Anatomical labels were identified using the SPM Anatomy Toolbox. Results were obtained through generalized
psycho-physiological interaction (gPPI) analysis (minimum voxel extent of 20 voxels and p <  0.001 significance
threshold, uncorrected).

Methods
Stimuli Selection.  In a pre-experiment survey, three photographs of an actress expressing a basic emo-
tional valence (positive, ambiguous and negative) were chosen as emotional cues for the current study (Fig. 1b).
“Ambiguous” was defined as a neither positive nor negative rating in valence and arousal44. These stimuli were
first developed for a previous, behavioral study of emotional improvisation13, and were intended to represent an
emotion with minimal distractions, and without eliciting a strong emotional reaction from perceivers. Photos
showed an actress from the collarbone upward, looking away from the camera, and photos were desaturated so
there would be no color cues. We developed these visual cues for emotions in order to avoid potential confounds
of linguistic labels45.

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Figure 5.  PPI Results. (A) Regions showing significantly higher (in red) effective connectivity with
the left amygdala (x = −22, y = −2, z = −22) during positive compared to negative improvisation
[PosImprov > NegImprov contrast in gPPI analysis]. (B) Regions showing significantly higher (in red) or
lower (in blue) effective connectivity with the left insula (x =  − 43, y =  14, z =  − 4) during positive compared
to negative improvisation [PosImprov >  NegImprov contrast in gPPI analysis]. Contrasts are thresholded at
p <  0.001 (uncorrected) with a minimum cluster size of 20 voxels. Panels are plotted in the sagittal plane, with
the x-coordinate of each panel indicated in the lower left-hand corner of each slice. Tick marks on the y-axis of
each panel indicate differences of 10 mm in the y dimension of the image space. Tick marks on the x-axis of each
panel indicate differences of 10 mm in the z dimension of the image space. Dashed lines through the center of
each slice converge on the origin of the image in the axial and coronal planes. All coordinates are presented in
MNI space.

11 males and 9 females (mean age =  32 ±  =  17s.d.), from the Johns Hopkins University community rated a
selection of images on a visual analog scales based on Russell’s circumplex model36,44, and results were coded on
a nine point scale (0–9, Negative-Positive). We calculated a one-way ANOVA with factors Emotion (Negative,
Ambiguous, Positive). Tukey’s honestly significant difference criterion was used for post hoc comparisons. A sig-
nificant main effect of Emotion [F(1, 2) =  110.87, p <  0.001] was observed. Mean ratings for the stimuli: Negative,
mean =  2.95, s.d. =  1.09, Ambiguous, mean =  4.3, s.d. =  0.80, Positive, mean =  7.5, s.d. =  1.05. A full description of
our stimuli pre-testing is available in McPherson et al.13. Informed consent was obtained in writing from all subjects,
and all experimental procedures were approved by the Johns Hopkins University School of Medicine Institutional
Review Board. All experimental procedures were carried out in accordance with the approved guidelines.

Musical Performance Analysis.  We analyzed the MIDI (Musical Instrument Digital Interface) piano out-
put obtained during fMRI scanning using measures of salient musical features including note density, note dura-
tion distribution, note maxima and minima, mode, and key. These measures were compared for the chromatic
scales and improvisations created in response to the different emotional targets. These results were calculated
using the MIDI Toolbox46, and a complete explanation of the calculation of these features can be found in Eerola
and Toiviainen, 2004.
Note density is a measure of notes per second, and for monophonic compositions can be used as an indication
of tempo (higher note densities generally correspond with faster tempos). Note maxima and minima indicate the
highest and lowest pitch, respectively, played in a given musical segment. For note density, maxima and minima,
we calculated a one-way ANOVA with the within-subject factor Emotion (Negative, Ambiguous, Positive) for
both improvisation and chromatic scale trials. Tukey’s honestly significant difference criterion was used for post
hoc comparisons.
The duration distribution function of the MIDI Toolbox returns the percentage of notes that fall into nine dif-
ferent logarithmically organized bins (note length categories). Length categories are defined as a unit of beats. We
set our MIDI tempo so that 1 beat =  0.5 s (quarter note =  120 Beats Per Minute (BPM)). Therefore, bin 1 =  1/8 s,
bin 3 =  ¼ s, bin 5 =  ½ s, bin 7 =  1 s, and bin 9 =  2 s. The relationship between bin 1 and bin 9 is proportional to the
relationship between a sixteenth note and a whole note. We compared corresponding duration distribution bins
using two-sample Kolmogorov-Smirnov tests.

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Key (tonal center) and mode (major vs. minor) were calculated using the Krumhansl & Schmuckler (K-S)
key-finding algorithm, which uses the pitch class distribution of a piece (weighted according to duration) to return
a key profile for the piece46. We used the K-S key finding algorithm to determine the best fit for each entire 44 s
improvisation. Mode and key calculations were confirmed by the authors through a visual inspection of the scores.

Functional Neuroimaging Testing.  Subjects.  Twelve professional jazz pianists (11 male, 1 female; mean
age =  39.9 ±  15.8) participated in the study. All subjects had been performing piano professionally for over 5
years (mean yrs performing professionally =  18.35 ±  13.28). Subjects were recruited as they became available,
without an a-priori regard to balancing by gender. None of the subjects reported histories of neurologic, auditory
or psychiatric disorders. Informed consent was obtained in writing for all subjects, and the research protocol was
approved by the Johns Hopkins School of Medicine Institutional Review Board.

Experimental Design.  A block-design imaging paradigm was used to assess the effect of emotional intent on
musical creativity (Fig. 1a). Rest blocks were 16 seconds in duration, and test blocks were 44 seconds in duration.
While in the scanner, the pianists were shown the three selected photographs of an actress representing, ‘Positive’,
‘Negative’, and ‘Ambiguous’ emotions. At the beginning of the presentation of each image, subjects were given a
simultaneous matching visual and auditory cue instructing them to respond in a specific way to the image. This cue
lasted three seconds. The cues were: “View”, “Chromatic Scale” and “Improvise”. Subjects were instructed to simply
fixate on the image and keep their eyes open during the entire ‘View’ condition. The chromatic scale condition was
designed to assess neural activity during a highly constrained, non-creative and non-emotional musical motor
task. For the chromatic condition, subjects were told to play an ascending and descending chromatic scale over
the entire range of the keyboard. Pianists were instructed to make this chromatic scale the same tempo regard-
less of the picture they were viewing. Before scanning began, pianists were familiarized with the target tempo,
approximately eighth note =  180 BPM (three notes per second), and were instructed to keep this tempo consistent
between blocks. For the improvise condition, subjects were instructed to improvise a composition that they felt
best represented the emotion expressed by the images. Improvisation was unrestricted melodically, harmonically,
and rhythmically, but the subjects were instructed to play monophonically (one note at a time) using their right
hand. Pianists were restricted to using their right hand while improvising and playing chromatic scales due to
space considerations on the scanner piano. Stimuli were presented in a pseudorandom order, and there were 24
test blocks per subject, with 8 per emotion (4 Improvisation, 2 Chromatic and 2 View). Subjects were asked to
keep their eyes open during the entire experiment, even rest blocks, and to refrain from moving their head or any
other part of their body other than their right hand.

Procedure.  During scanning, subjects used a custom-built non-ferromagnetic piano keyboard (MagDesign,
Redwood, CA) with thirty-five full-size plastic piano keys. The piano keyboard was placed on the subject's lap in
supine position, while their knees were elevated with a bolster. A soft velcro square was placed on Middle C of the
piano, allowing subjects to orient their hand without viewing the keyboard. Subjects were visually monitored to
ensure that they did not move their left (non-playing) hand, head, trunk, or other extremities during performance.
Before the experiment commenced, subjects were given time to find a comfortable playing position and familiarize
themselves with the keyboard and environment. Subjects also had a trial run to test sound levels, make final key-
board placement adjustments, and practice the paradigm before scanning began. Data from these test blocks were
discarded, and the test block was repeated if the subject requested more time to become comfortable in the scanner.
The scanner keyboard had MIDI output, which was sent to a Macintosh MacBook Pro laptop computer running
the Logic Express 8 sequencing environment (Apple Inc., Cupertino, CA). Piano sound output was routed back to
the subject via in-ear electrostatic earspeakers (Stax, Saitama, Japan). In addition to the electrostatic earspeakers,
subjects wore additional ear protection to minimize background scanner noise. For each subject, ear speaker
volume was set to a comfortable listening level that could be easily heard over the background scanner noise. A
double mirror mounted on the head coil above the subject's eyes allowed them to view a rear projection screen
behind the scanner bore. The stimuli and instructions were presented with EPrime47.

Image acquisition.  All scans were performed at the F.M. Kirby Research Center for Functional Brain Imaging
at the Kennedy Krieger Institute of Johns Hopkins University. Blood oxygen level dependent imaging (BOLD)
data and T1-weighted anatomical images were acquired using a 3-Tesla whole-body scanner (Philips Electronics,
Andover, MA) using an sixteen-channel head coil and a gradient-EPI sequence. The following scan parameters
were used: TR =  2000 ms, TE =  30 ms, flip-angle =  75 degrees, field of view 216.000 ×  128.000 ×  240.000 mm, 32
parallel axial slices covering the whole brain, 4 mm thickness (3 ×  3 mm in-plane resolution). 720 volumes were
acquired for each subject.

fMRI Analysis.  Standard preprocessing steps were completed in SPM8, including realignment to the first vol-
ume of the run, coregistration with a participant's T1-weighted structural image, indirect normalization of the
structural image to template space, propagation of normalization parameters to coregistered functional images,
and smoothing with an 8 mm FWHM kernel. A first-level general linear model was estimated for each sub-
ject using ten regressors, one for rest and one for each experimental condition combination—emotion (posi-
tive, negative, ambiguous—Pos, Neg and Amb) and task (view, chromatic scale, improvisation—View, Chrom
and Improv). Each regressor was convolved with a standard hemodynamic response function. Design matri-
ces also included covariates of non-interest, which consisted of motion parameters calculated during the rea-
lignment stage and mean signal intensity for the run. Between-emotion (e.g. [PosImprov >  NegImprov]) and
within-emotion (e.g. [PosImprov >  PosChrom]) contrasts were estimated for each subject. Contrasts were then
entered into a second-level random-effects model using a one-sample t-test. Random-effects analyses take into

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account inter-subject variability, and therefore can be generalized to a broader population. Contrasts were thresh-
olded at an uncorrected p value of 0.005 with a minimum voxel extent of 10 voxels. Analysis of average effect sizes
was completed using the rfxplot toolbox48.

PPI Analysis.  Psycho-physiological interaction (PPI) analysis can be used to identify task-dependent changes
in effective connectivity between a seed region and other regions in the brain49. We used the generalized
psycho-physiological interaction (gPPI) toolbox50 to examine differences in the networks of brain areas that exhib-
ited functional connectivity with emotion-specific brain areas during improvisation conditions with different
emotional intention. Seed regions for this analysis were derived from emotional condition contrasts estimated in the
primary analysis. Significant shifts in functional connectivity in an emotional condition for each seed region were
identified by applying inclusive masking for within-emotion (i.e. [PosImprov >  PosChrom]) and between-emotion
(i.e. [PosImprov >  NegImprov]) contrasts, with a minimum voxel extent of 20 voxels and p <  0.001 significance
threshold, uncorrected.

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Acknowledgements
This project was funded by the Dana Foundation and the Brain Science Institute of Johns Hopkins University School
of Medicine, as well as a Training Grant (T32-DC000023) from the Department of Biomedical Engineering, Johns
Hopkins University. The funders had no role in study design, data collection and analysis, decision to publish, or
preparation of the manuscript.

Author Contributions
M.J.M. designed and performed experiment, analyzed data and wrote the manuscript. F.S.B. analyzed data and
wrote the manuscript. M.L.G. designed and performed experiment. P.J. performed experiment. C.J.L. supervised
the project and wrote the manuscript.

Additional Information
Competing financial interests: The authors declare no competing financial interests.
How to cite this article: McPherson, M. J. et al. Emotional Intent Modulates The Neural Substrates Of
Creativity: An fMRI Study of Emotionally Targeted Improvisation in Jazz Musicians. Sci. Rep. 6, 18460;
doi: 10.1038/srep18460 (2016).
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Scientific Reports | 6:18460 | DOI: 10.1038/srep18460 14