A Systematic Review on the Neural Effects of Music on ...A Systematic Review on the Neural Effects of Music on Emotion Regulation: Implications for Music Therapy Practice Kimberly

A Systematic Review on the NeuralEffects of Music on Emotion Regulation:Implications for Music Therapy Practice

Kimberly Sena Moore, MM

University of Missouri-Kansas City

Background: Emotion regulation (ER) is an internal process throughwhich a person maintains a comfortable state of arousal by modulatingone or more aspects of emotion. The neural correlates underlying ERsuggest an interplay between cognitive control areas and areas involved inemotional reactivity. Although some studies have suggested that musicmay be a useful tool in ER, few studies have examined the links betweenmusic perception/production and the neural mechanisms that underlie ERand resulting implications for clinical music therapy treatment. Objectivesof this systematic review were to explore and synthesize what is knownabout how music and music experiences impact neural structuresimplicated in ER, and to consider clinical implications of these findingsfor structuring music stimuli to facilitate ER.

Methods: A comprehensive electronic database search resulted in 50studies that met predetermined inclusion and exclusion criteria.Pertinent data related to the objective were extracted and studyoutcomes were analyzed and compared for trends and commonfindings.

Results: Results indicated there are certain music characteristics andexperiences that produce desired and undesired neural activationpatterns implicated in ER. Desired activation patterns occurred whenlistening to preferred and familiar music, when singing, and (in musicians)when improvising; undesired activation patterns arose when introducingcomplexity, dissonance, and unexpected musical events. Furthermore,

Kimberly Sena Moore, MM, Conservatory of Music and Dance, University ofMissouri-Kansas City.

This research was supported in part by a Graduate Assistance Fund award fromthe University of Missouri-Kansas City Women’s Council. The author would like tothank Mirna Herrera and Brittany Slaughter for their assistance as coders, and Dr.Deanna Hanson-Abromeit for her guidance and mentorship with this project. Thisproject was completed in partial fulfillment of a doctoral degree.

Address correspondence concerning this article to Kimberly Sena Moore,Conservatory of Music and Dance, University of Missouri-Kansas City, Kansas City,MO 64119-2229. Email: [email protected]

Journal of Music Therapy, 50(3), 2013, 198–242G 2013 by the American Music Therapy Association

the connection between music-influenced changes in attention and itslink to ER was explored.Conclusions: Implications for music therapy practice are discussed andpreliminary guidelines for how to use music to facilitate ER are shared.

Keywords: music; emotion regulation; neuroscience, amygdala

Self-regulation is a complex process of self-directed change. It is theability to implement processes and actions that allow one to effectivelycontrol and manage multiple levels of experiences: cognitive,emotional, behavioral, and physiological (Bandura, 1991; Blaustein& Kinniburgh, 2010; Karoly, 1993; Larsen, 2000; Smith-Donal, Raver,Hayes, & Richardson, 2007). Self-regulation is considered a centraldevelopmental milestone in early childhood (Liebermann, Gies-brecht, & Muller, 2007) that has lifelong implications for one’semotional, cognitive, social, and mental health. An essentialcomponent of self-regulation is emotion regulation (ER) (Diamond& Aspinwall, 2003; Geva & Feldman, 2008) and the development ofER is considered an early marker for the development of appropriateself-regulation (Cole, Dennis, Smith-Simon, & Cohen, 2009).

There has been a significant increase in the past 25 years inexploring the neural correlates underlying a phenomenon; ER isno exception. In addition, there has been increased interest in themusic neurosciences and in studying the neural correlatesunderlying music perception and cognition. Although it has longbeen considered that music influences emotions, there has beenlittle attention given to understanding how music affects emotionregulation. For the music therapist, an understanding of thisphenomenon has therapeutic implications for a variety of clinicalpopulations that find it a challenge to control and manage theiremotional experiences. Thus, the major goal of this review was toexplore what is known about how music and music experiencesimpact neural structures implicated in emotion regulation, and toconsider possible clinical implications of these findings.

Emotion Regulation

Emotion regulation is an internal process through which aperson is able to maintain a comfortable state of arousal bymodulating one or more aspects of emotion (Blaustein &Kinniburgh, 2010; Diamond & Aspinwall, 2003; McRae et al.,

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2010). It involves using strategies and processes designed to create anew emotional response or change a current one (Gyurak, Gross, &Etkin, 2011; McRae et al., 2010; Ochsner & Gross, 2005) that can beexplicit (e.g., effortful or conscious) or implicit (e.g., automatic orunconscious) (Diamond & Aspinwall, 2003; Gyurak et al., 2011).Generally, successful emotion regulation strategies either alter theway an individual attends to a situation, interprets the meaning ofa situation (McRae et al., 2010), or changes the situation itself(Diamond & Aspinwall, 2003). Difficulties with emotion regulationcan have a life-long impact on an individual’s mental health andwell-being (Saxena, Dubey, & Pandey, 2011). Many researchersconsider appropriate emotion regulation to be a marker of mentalhealth as it allows a person flexibility in how he or she responds andreacts to situations and moments of distress (Cole et al., 2009;Gyurak et al., 2011; McRae et al., 2010). There are many disordersand syndromes in which difficulties with emotion regulation can bea challenge, including Attention Deficit/Hyperactivity disorder,Autism and Asperger syndrome (Masao, 2004), Post-TraumaticStress Disorder (PTSD), and trauma (Ehring & Quack, 2010).

The neural correlates underlying emotion regulation suggest aninterplay between frontal lobe areas involved in cognitive controland areas involved in emotional reactivity (Gyurak et al., 2011).More specifically, cognitive control areas include the lateralprefrontal cortex (Gyurak et al., 2011; McRae et al., 2010; Ochsner& Gross, 2005), the orbitofrontal cortex (Masao, 2004; Ochsner &Gross, 2005; Rempel-Clower, 2007; Schore, 2001), and the anteriorcingulate cortex (Gyurak et al., 2011; McRae et al., 2010; Ochsner &Gross, 2005). The amygdala is the primary structure implicated inemotional reactivity (Gyurak et al., 2011; Masao, 2004; McRae et al.,2010; Ochsner & Gross, 2005). In general, emotion regulation ischaracterized by increased activation in the cognitive control andmonitoring areas—the anterior cingulate cortex, orbitofrontalcortex, and lateral prefrontal cortex—which leads to decreasedactivation in the amygdala (Gyurak et al., 2011; Ochsner & Gross,2005; McRae et al., 2010; Rempel-Clower, 2007).

Music and Emotion Regulation

Music has long been thought to influence emotions andemotion control. In his seminal book The Anthropology of Music,Merriam (1964) wrote about music’s role as a producer of

200 Journal of Music Therapy

emotions. It has also been noted that music can evoke emotions inlisteners (Sloboda & Juslin, 2010) and may be an effective moodinduction technique (Thaut & Wheeler, 2010). However, moodinduction is different than emotion regulation; moods areaffective states lower in intensity than emotions (Juslin & Sloboda,2010). Furthermore, evoking emotions is a general conceptwhereas emotion regulation is geared towards the specific goalof maintaining a comfortable state of arousal (Blaustein &Kinniburgh, 2010; Diamond & Aspinwall, 2003; McRae et al.,2010). More recent research has focused on the neural basisunderlying music-evoked emotions, finding that music doesindeed impact neural areas implicated in emotion processing(Blood & Zatorre, 2001; Koelsch, 2010; Trainor & Schmidt, 2003).However, much in the literature is from the music neurosciencefield and focuses on the neural mechanisms underlying musiclistening, playing, or improvisation. There is little that exploresthe connection between music processing and clinical treatment;as such, there is little in the way of clinical implications relevant tothe music therapy clinician. Therefore, the purpose of thisexploratory review was to synthesize findings from studies thatreported on the effect of music and music-based experiences onneural structures implicated in ER, and to create preliminaryclinical considerations based on this synthesis.

Description of the Condition

For the purposes of this review, ER was defined as an internalprocess through which a person is able to maintain a comfortablestate of arousal by modulating one or more aspects of emotion(Blaustein & Kinniburgh, 2010; Diamond & Aspinwall, 2003; McRaeet al., 2010). It is characterized by the involvement of the amygdala(Gyurak et al., 2011; Masao, 2004; McRae et al., 2010; Ochsner &Gross, 2005), the anterior cingulate cortex (Gyurak et al., 2011;McRae et al., 2010; Ochsner & Gross, 2005), the orbitofrontalcortex (Masao, 2004; Ochsner & Gross, 2005; Rempel-Clower, 2007;Schore, 2001), and the lateral prefrontal cortex (Gyurak et al., 2011;McRae et al., 2010; Ochsner & Gross, 2005). There are cases where‘‘emotion regulation’’ has the same meaning as ‘‘affect regulation’’(Schore, 2001), although there are times when affect regulationrefers to a set of intervention techniques (Verheugt-Pleiter, 2008);the latter did not fit the intent of this review.

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Description of the Stimulus

For the purposes of this review, ‘‘music’’ referred to any acousticstimulation provided by a complex, organized sound. As long as thestudy fit other criteria for this review, ‘‘music’’ also included musicalproperties, such as rhythm, musical interval, harmony, and pitch,and music experiences, such as listening to music, playing aninstrument, improvising, or composing. The literature lists ‘‘music’’most often, with occasional references to terms such as ‘‘acousticstimulation,’’ ‘‘music therapy,’’ and ‘‘complex musical soundstimuli.’’ Research using rhythm that was associated with circadianrhythm, cardiac rhythm, respiratory rhythm, dietary rhythm, andother nonmusical references to rhythm were excluded.

Objectives

1. To explore and synthesize results examining the effects ofmusic on neural structures implicated in emotion regulation.

2. To create preliminary clinical considerations for structuringthe music stimulus when facilitating emotion regulation.

Methods

Search Strategies

The search and analysis processes used in this review wereconsistent with those outlined by Cooper (1998) and Khan, Kunz,Kleijnen, and Antes (2011). Studies considered for this reviewwere published through April 2012 and identified through acomprehensive search in the following electronic databases:MEDLINE, PsycINFO, CINAHL, SIGLE, National Institute forHealth Research, Current Controlled Trials, ClinicalTrials.gov,and CAIRSS for Music. Electronic databases were searched usingthe following keyword phrases: ‘‘music and amygdala,’’ ‘‘musicand orbitofrontal cortex,’’ ‘‘music and anterior cingulate,’’ and‘‘music and prefrontal.’’ Search results generated from the ‘‘musicand prefrontal’’ keyword phrase were scanned and included forconsideration if they included the words ‘‘dorsolateral,’’ ‘‘ventro-lateral,’’ or ‘‘lateral.’’

Article Inclusion Criteria

1. The article was a primary research study.


2. Participants were typically-developing humans, with norestrictions as to age, gender, ethnicity, or type of setting.

3. Music was the primary stimulus, regardless of how it wasimplemented (e.g., singing, listening, improvising, etc.), thegenre of music, or the music instrument(s) incorporated.

4. Study results reported on the impact of music on one ormore of the following neural structures: amygdala, anteriorcingulate cortex, orbitofrontal cortex, and lateral prefrontalcortex.

5. Articles were published in English.6. Articles were published in peer-reviewed journals.

Article Exclusion Criteria

1. The article was a review study or theoretical paper.2. Participants had a disorder, brain damage (such as lesions

and excisions), or a syndrome.3. The study of rhythm was associated with circadian rhythm,

cardiac rhythm, respiratory rhythm, dietary rhythm, andother nonmusical references to rhythm.

Data Collection and Analysis Process

The author and two research assistants independently extractedpertinent information from the included studies related to thefirst objective. This included, as applicable, information aboutparticipant characteristics (i.e., number of participants, sexdistribution, average age, age range, and musical ability), studydesign (i.e., primary research question, neural measurement toolutilized, and other behavioral and neuropsychological toolsutilized), characteristics of the music and/or music experienceutilized (i.e., type of music experience, music characteristicstudied, music instrument used, and genre of musical stimulus),and study outcomes (i.e., general outcomes of the study andneural structure-specific outcomes). Pertinent data related to theobjective were extracted and study outcomes were analyzed andcompared for trends and common findings. Differences in dataextracted were discussed and results agreed upon for dataanalysis. Initial interrater reliability among the three coders was75.5%. When controlled for typographical errors and updates tothe coding sheet that occurred as a result of discussing differencesin data extracted, final inter-rater reliability was 83.6%.

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Results of the Search

The comprehensive electronic database search resulted in 319articles that were evaluated for inclusion according to theinclusion and exclusion criteria listed above. Of those initialstudies, 103 met the criteria. Forty-four (44) studies wereduplicates, which led to the inclusion of 59 unique studies.During the coding process, an additional nine studies wereexcluded for the following reasons: the neural structures were notactivated as a result of music stimulation (Dehaene-Lambertz etal., 2010; Schulze, Zysset, Mueller, Friederici, & Koelsch, 2011;Sluming et al., 2002), music was not used as the intervention(Blasi et al., 2011; Haslinger et al., 2004; Milton, Solodkin,Hlustik, & Small, 2007), the study was a dissertation (Chapin,2010), the study offered preliminary data that was reported in asecond study (Zarate & Zatorre, 2005), and the study did notreport information clearly enough for data extraction (Bodner,Muftuler, Nalciogly, & Shaw, 2001). Thus, this review included 50research studies.

Results

Characteristics of Included Studies

Participant characteristics. There were a total of 811 partici-pants in the studies (M 5 25.1 years; SD 5 6.2 years, range: 12–60 years), 757 adults and 54 adolescents. Over half of theparticipants were male (54.7%) and the rest were female (45.3%).Studies used an average of 16 participants (range: 6–49participants). Almost half of the studies used musicians asparticipants or a combination of musicians and nonmusicians(44.4%) and the remainder either used nonmusicians or authorswere not specific in the reporting (Table 1).

Characteristics of study design. Frequency information, relatedto study design characteristics of the included studies, is reportedin Table 1. The two most common neural measurement toolsused were functional Magnetic Resonance Imaging (fMRI)(65.5%) and Positron Emission Tomography (PET) (18.2%).Other techniques used included electroencephalography(EEG), event-related potentials (ERPs), magnetoencephalogra-phy (MEG), and a brain oxygen measurement tool called an OTsystem. Studies included in this review incorporated a variety of


musical experiences. The majority of studies (69.6%) usedlistening to recorded music as the experience, followed bysinging, instrument playing, and improvisation. The mostcommon musical instrument used, when specified, was piano(41.7%) followed by voice (18.3%), and the most commonmusical genre used, when specified, was tonal, Western instru-mental music (43.1%), followed by popular or current music, jazzor improvisation, and nonwestern music. Almost half of the time(49.1%) researchers were not investigating a particular musicalelement (i.e., rhythm, pitch, harmony, etc.). If they did, harmonywas investigated the most frequently (21.8%), followed by rhythm,melody, pitch, timbre, and pitch interval.

Synthesis of Results

A summary of pertinent characteristics and outcome measures ofindividual studies is reported in Table 2 and a synthesis of the mainfindings is reported in Table 3. Anterior cingulate cortex activationwas reported and/or described the most frequently (32.9%),followed by lateral prefrontal cortex activation, amygdala activation,and orbitofrontal cortex activation (Table 1). Results will bepresented in a bottom-up fashion, from the deepest neural structure,the amygdala, to the most superficial area, the prefrontal cortex.

Amygdala. Multiple studies reported that the amygdala wasactivated when listening to minor, dissonant, negative, orunpleasant music (Koelsch, Fritz, Cramon, Muller, & Friederici,2006; Lerner, Papo, Zhdanov, Belozersky, & Hendler, 2009;Pallesen et al., 2005). Amygdala activation occurred during anunexpected event (e.g., hearing an irregular chord) and itsactivity could be modulated by a single chord change (Koelsch,Fritz, & Schlaug, 2008). One study reported that the amygdala wasactivated during music listening regardless of consonance ordissonance (Ball et al., 2007). Amygdala activation increased whenlistening to music with eyes closed (Lerner et al., 2009) and wasactivated more strongly when music was paired with visual stimulias compared to no visual stimuli (Baumgartner, Lutz, Schmidt, &Jancke, 2006; Eldar, Ganor, Admon, Bleich, & Hendler, 2007).One study noted a lateralization effect, reporting that the rightamygdala was activated more strongly during an audiovisualcondition than the left amygdala, and its activity increased withexposure over time (Dyck et al., 2011). The right amygdala in

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TABLE 1Frequency of Participant and Study Characteristics

Category Number Relative frequency

Music training of participants1. Nonmusicians only 15 30%1. Musicians only 13 26%1. Not specified 13 26%1. Both musicians and nonmusicians 9 18%

Neural measurement tools used1. fMRI 36 65.5%1. PET 10 18.2%1. Electroencephalography (EEG) 5 9.1%1. Event-related potentials (ERP) 2 3.6%1. Magnetoencephalography (MEG) 1 1.8%1. Event-related potentials (ERP) 1 1.8%

Type of music experience1. Music listening (recorded music) 39 69.6%1. Singing 8 14.3%1. Instrument playing 5 8.9%1. Improvisation 4 7.1%

Music instrument(s) utilized1. Piano 25 41.7%1. Voice 11 18.3%1. Instrumental (orchestral) 8 13.3%1. Instrumental (electronic) 5 8.3%1. Not specified 4 6.7%1. Instrumental (accompaniment) 3 5.0%1. Instrumental (solo) 2 3.3%1. Instrumental (nonwestern) 1 1.7%1. Other 1 1.7%

Genre of music1. Western tonal instrumental music 22 43.1%1. Not specified 13 25.5%1. Popular/current music 6 11.8%1. Jazz/improvisation 4 7.8%1. Nonwestern music 3 5.9%1. Western tonal vocal music 2 3.9%1. Other 1 2.0%

Music element studied1. Not specified 27 49.1%1. Harmony 12 21.8%1. Other 6 10.9%1. Rhythm 3 5.5%1. Melody 2 3.6%


particular was recruited during ‘‘sad’’ music (Mitterschiffthaler,Fu, Dalton, Andrew, & Williams, 2007), is implicated in earlyneural responses to chord violations (James, Britz, Vuilleumier,Hauert, & Michel, 2008), and exhibited chord-dependentresponses (Pallesen, Brattico, Bailey, Korvenoja, & Gjedde,2009). The left amygdala had consistently similar activationpatterns, with increased activation reported when listening tomusic rated with a higher negative emotional valence (Dyck et al.,2011). The amygdala was deactivated during music improvisation(Limb & Braun, 2008) and when listening to pleasant music(Blood & Zatorre, 2001; Koelsch et al., 2006). A long-termhabituation effect was noted, perhaps due to the amygdala’s rolein evaluating salience (Mutshuler et al., 2010). To summarize,amygdala activation and deactivation patterns changed based onthe type of music experience, the characteristics of the musicstimulus, and perceived valence of the music.

Anterior cingulate cortex (ACC). The ACC was activated duringvoluntary pitch correction (Zarate, Wood, & Zatorre, 2010),discrimination tasks (Brown & Martinez, 2007), when listening tochord violations (James et al., 2008), and when monitoringperformance errors (Ruiz, Jabusch, & Altenmuller, 2009). Inaddition, listening to familiar music activated the ACC (Janata,2009) as did, in musicians, listening to dissonant chords (Foss,Altschuler, & James, 2007). ACC activation increased during bothsinging (Perry et al., 1999) and music listening tasks (Menon &

Category Number Relative frequency

1. Pitch 2 3.6%1. Timbre 2 3.6%1. Interval 1 1.8%

Neural structure reported on1. Anterior cingulate cortex (ACC) 24 32.9%1. Lateral prefrontal cortex (lateral PFC) 21 28.7%1. Amygdala 12 16.4%1. Orbitofrontal (OFC) 9 12.3%

Note. Some studies incorporated multiple types of neural measurement tools, musicexperiences, music instruments, music genres, studied multiple music elements,and reported on multiple neural structures. These numbers are reflected in thereported frequencies.

TABLE 1Continued

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li;(2

)H

eart

rate

and

resp

irat

ion

incr

ease

dw

hile

liste

nin

gto

mu

sic

that

elic

ited

chill

s.B

row

n&

Mar

tin

ez(2

006)

fMR

Ist

ud

yex

plo

rin

gn

eura

lm

ech

anis

ms

un

der

lyin

gm

elo

dy

and

har

mo

ny

dis

crim

inat

ion

11M

usi

cian

sM

usi

cli

sten

ing

(rec

ord

ed)

Har

mo

ny,

Mel

od

yA

CC

,d

lPF

C(1

)D

iscr

imin

atio

np

roce

ssin

gin

volv

esd

om

ain

-sp

ecif

icse

nso

rim

oto

rar

eaan

dd

om

ain

-gen

eral

wo

rkin

gm

emo

ryan

der

ror

det

ecti

on

area

s;(2

)Si

mil

arac

tiva

tio

np

atte

rns

bet

wee

nm

elo

dy/

har

mo

ny

pro

cess

ing

and

bet

wee

np

erce

pti

on

/p

rod

uct

ion

ofm

usi

c,in

clu

din

gd

lPF

Can

dA

CC

Bro

wn

etal

.(2

004)

PE

Tst

ud

yex

plo

rin

gn

eura

lm

ech

anis

ms

un

der

lyin

gp

leas

ant

feel

ings

10N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)N

ot

spec

ifie

dA

CC

(1)

Spon

tan

eou

sac

tivat

ion

oflim

bic

and

par

alim

bic

area

sd

uri

ng

task

-fr

ee,p

assi

velis

ten

ing

toun

fam

iliar

but

liked

mus

ic;(

2)St

ron

ger

left

-hem

isp

her

eac

tivat

ion

with

mu

sic-

elic

ited

pos

itive

emot

ion

s

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Bro

wn

etal

.(2

006)

PE

Tst

ud

yex

plo

rin

gn

eura

lm

ech

anis

ms

un

der

lyin

gm

elo

dic

and

spee

chge

ner

atio

n

10M

usi

cian

sM

usi

cli

sten

ing

(rec

ord

ed),

Sin

gin

g

Mel

od

yA

CC

(1)

Sign

ific

ant

ove

rlap

inst

ruct

ure

sac

tiva

ted

du

rin

gm

elo

dic

imp

rovi

sati

on

and

sen

ten

cege

ner

atio

n,

incl

ud

ing

AC

C;

(2)

Som

ela

tera

liza

tio

nd

iffe

ren

ces,

such

asst

ron

ger

left

-h

emis

ph

ere

acti

vati

on

wit

hse

nte

nce

gen

erat

ion

Cal

lan

etal

.(2

006)

fMR

Ist

ud

yin

vest

igat

ing

the

neu

ral

dif

fere

nce

sb

etw

een

per

ceiv

ing

and

pro

du

cin

gso

ng

and

spee

ch

16N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

),si

ngi

ng

No

tsp

ecif

ied

OF

CD

iffu

se,

bil

ater

aln

etw

ork

of

ove

rlap

pin

gn

eura

lp

roce

sses

un

der

lie

per

cep

tio

nan

dp

rod

uct

ion

of

spee

chan

dso

ng,

sup

po

rtin

gre

lati

on

ship

bet

wee

nth

emC

oen

etal

.(2

009)

fMR

Ist

ud

yex

plo

rin

gef

fect

of

neg

ativ

eem

oti

on

alst

imu

lio

nn

eura

lp

roce

ssin

g

12N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

Not

spec

ified

AC

C,d

lPFC

(1)

Evi

den

ceo

fri

ght

hem

isp

her

ed

om

inan

cew

hen

pro

cess

ing

neg

ativ

eem

oti

on

s;(2

)R

igh

tin

sula

and

righ

tA

CC

seem

inte

gral

toaw

aren

ess

of

emo

tio

n;

(3)

Sad

nes

sp

erce

pti

on

incr

ease

dw

hen

list

enin

gto

sad

mu

sic

bu

td

idn

ot

affe

ctp

ain

per

cep

tio

n

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 211

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

de

Man

zan

o&

Ull

en(2

012)

fMR

Ist

ud

yex

plo

rin

gn

eura

lm

ech

anis

ms

un

der

lyin

gre

spo

nse

gen

erat

ion

18M

usi

cian

sIm

pro

visa

tio

n,

inst

rum

ent

pla

yN

otsp

ecifi

edA

CC

,dlP

FC(1

)Si

gnif

ican

to

verl

apin

acti

vati

on

du

rin

gim

pro

visa

tio

nan

dsi

ght-

read

ing,

sugg

esti

ng

thes

ere

gio

ns

fulf

ill

gen

eric

fun

ctio

ns

infr

eege

ner

atio

nre

gard

less

of

goal

;(2

)H

igh

erac

tivi

tyd

uri

ng

pse

ud

o-

gen

erat

ion

task

inat

ten

tio

n,

wo

rkin

gm

emo

ry,

and

exec

uti

veco

ntr

ol

area

sD

yck

etal

.(2

011)

fMR

Ist

ud

yex

plo

rin

gam

ygd

ala

resp

on

sew

hen

pre

sen

ted

wit

hvi

sual

v.au

dio

visu

alin

pu

t

30N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

Am

ygd

ala

(1)

Am

ygd

ala

may

be

imp

lica

ted

inem

oti

on

regu

lati

on

,no

tju

stem

oti

on

per

cep

tio

n;

(2)

Rep

ort

edle

ft-la

tera

lize

dco

gnit

ive/

inte

nti

on

alco

ntr

ol

of

mo

od

and

righ

t-la

tera

lize

dau

tom

atic

ind

uct

ion

of

emo

tio

n;

(3)

Rep

ort

edst

ron

ger

acti

vati

on

wit

hau

dio

visu

alin

pu

tE

ldar

etal

.(2

007)

fMR

Ist

ud

yex

plo

rin

gli

mb

icre

spo

nse

sto

mu

sic

wh

enp

aire

d/

no

tp

aire

din

are

al-

wo

rld

con

text

12N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

Am

ygd

ala,

LP

FC(1

)B

rain

may

hav

ep

refe

ren

tial

re-

spon

seto

emot

ion

alst

imu

liw

hen

asso

ciat

edw

itha

con

cret

eco

n-

text

;(2)

Mu

sic

can

elic

itgr

eate

rem

otio

nal

effe

ctw

hen

pai

red

with

aco

ncr

ete

visu

alst

imu

liv.

wh

enp

rese

nte

don

itsow

n

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Flo

res-

Gu

tier

rez

etal

.(2

007)

fMR

Ian

dE

EG

stu

dy

exp

lori

ng

neu

ral

acti

vity

asso

ciat

edw

ith

ple

asan

t/u

np

leas

ant

emo

tio

nal

stat

es

19N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)N

otsp

ecifi

edO

FC

(1)

Mu

sic-

elic

ited

emo

tio

ns

req

uir

eco

gnit

ive-

sen

sory

inte

grat

ion

and

hav

ew

ides

pre

adac

tiva

tio

no

fco

gnit

ive,

lan

guag

e,an

dem

oti

on

pro

cess

ing

area

s;(2

)D

iffe

ren

tst

ruct

ure

sin

volv

edin

pro

cess

ing

po

siti

vean

dn

egat

ive

mu

sica

lem

oti

on

sw

ith

left

sid

eac

tiva

ted

for

ple

asan

tem

oti

on

san

dri

ght

for

un

ple

asan

tF

ord

etal

.(2

011)

fMR

Ist

ud

yin

vest

igat

ing

neu

ral

mec

han

ism

su

nd

erly

ing

auto

bio

grap

hic

alm

emo

ries

asso

ciat

edw

ith

mu

sic

16N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

AC

C,L

PFC

(1)

Dif

fere

nt

neu

ral

stru

ctu

res

and

net

wo

rks

recr

uit

edfo

rd

iffe

ren

tty

pes

of

auto

bio

grap

hic

alm

emo

ry;

(2)

Sup

po

rts

use

of

mu

sic

asan

auto

bio

grap

hic

alre

trie

val

cue

for

mem

ori

esth

atar

ep

osi

tive

,em

oti

on

ally

char

ged

,an

dsu

bje

ctto

reli

vin

gas

itac

tiva

ted

area

sre

late

dto

emo

tio

np

roce

ssin

gan

dm

emo

ryre

trie

val

(e.g

.,V

PF

Can

dA

CC

)

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 213

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Fo

sset

al.

(200

7)fM

RI

stu

dy

exp

lori

ng

neu

ral

mec

han

ism

sim

pli

cate

din

Pyt

hag

ore

anra

tio

rule

s

13B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Mu

sic

list

enin

g(r

eco

rded

)In

terv

alA

CC

(1)

Neu

ral

acti

vati

on

of

dis

son

ant

inte

rval

ssi

gnif

ican

tly

grea

ter

than

con

son

ant

inte

rval

s;(2

)A

ctiv

atio

np

atte

rns

tod

isso

nan

ced

iffe

ren

tin

mu

sici

ans

(lef

t-lat

eral

ized

)v.

no

nm

usi

cian

s(r

igh

t-la

tera

lized

),su

gges

tin

gst

ron

ger

acti

vati

on

inm

usi

cian

s’la

ngu

age

pro

cess

ing

area

sF

uji

saw

a&

Co

ok

(201

1)fM

RI

stu

dy

exp

lori

ng

neu

ral

net

wo

rks

acti

vate

dw

hen

list

enin

gto

dif

fere

nt

Wes

tern

har

mo

nie

s

12N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

y3O

FC

,d

lPF

C(1

)R

epo

rted

acti

vati

on

pat

tern

sw

hen

pro

cess

ing

har

mo

nie

s;(2

)O

FCsi

gnif

ican

tly

acti

vate

dd

uri

ng

cho

rdch

ange

s;(3

)M

ovi

ng

fro

ma

ten

sio

nch

ord

toa

favo

red

cho

rdel

icit

sa

stro

ng

bra

inre

spo

nse

.G

reen

etal

.(2

008)

fMR

Ist

ud

yex

plo

rin

gn

eura

lre

spo

nse

sto

mu

sica

lm

od

es

21N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

yA

CC

(1)

Fou

nd

dif

fere

nti

alac

tiva

tio

no

fce

rtai

nst

ruct

ure

sw

hen

liste

nin

gto

min

or

v.m

ajo

rm

usi

c,w

hic

hm

ayb

ed

ue

tod

isso

nan

ceo

ro

vera

llq

ual

ity

of

sad

nes

s;(2

)M

ino

rm

elo

die

sac

tiva

ted

mo

relim

bic

area

sth

anm

ajo

rm

elo

die

s

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Hu

gdah

let

al.

(199

9)P

ET

scan

exp

lori

ng

acti

vati

on

pat

tern

sas

soci

ated

wit

hd

ich

oti

cli

sten

ing

12N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

Tim

bre

dlP

FC

(1)

Asy

mm

etry

effe

cts

wh

enp

rese

nti

ng

dif

fere

nt

stim

uli

inea

chea

r,w

ith

grea

ter

left

hem

isp

her

eac

tiva

tio

nfo

rsp

eech

and

grea

ter

righ

th

emis

ph

ere

acti

vati

on

for

mu

sic;

(2)

Stro

nge

rac

tiva

tio

nfo

rp

roce

ssin

gsp

eech

v.m

usi

c,p

erh

aps

bec

ause

aud

ito

ryco

rtex

spec

iali

zed

for

ph

on

olo

gica

lp

roce

ssin

g;(3

)d

lPF

Cm

ayp

lay

aro

lein

det

ecti

ng

com

ple

xti

mb

reJa

mes

etal

.(2

008)

ER

Pim

agin

gst

ud

yin

vest

igat

ing

neu

ral

dif

fere

nce

sm

usi

cal

trai

nin

gh

asin

har

mo

nic

pro

cess

ing

26B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

yrt

.Am

ygd

ala

(1)

Rap

id,

righ

t-la

tera

lize

dn

eura

lre

spo

nse

sto

cho

rdvi

ola

tio

ns

for

mu

sici

ans

v.n

on

mu

sici

ans,

sugg

esti

ng

that

mu

sic

trai

nin

gen

han

ces

righ

th

emis

ph

eric

do

min

ance

;(2

)A

myg

dal

aac

tiva

ted

wh

end

etec

tin

gh

arm

on

icin

con

gru

enci

es

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 215

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Jan

ata

(200

9)fM

RI

stu

dy

inve

stig

atin

gn

eura

lm

ech

anis

ms

un

der

lyin

gau

tob

iogr

aph

ical

mem

ori

esas

soci

ated

wit

hm

usi

can

dM

PF

Cac

tiva

tio

n

13N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

vlP

FC

(1)

mP

FC

asso

ciat

edw

ith

pro

cess

ing

mu

sic

and

mem

ori

es,

wit

ha

gen

eral

ized

acti

vati

on

incr

ease

bas

edo

nfa

mil

iari

tyan

dau

tob

iogr

aph

ical

sali

ence

of

mu

sic;

(2)

Res

ult

sd

emo

nst

rate

exte

nd

edau

tob

iogr

aph

ical

mem

ory

net

wo

rkth

atin

clu

des

mP

FC

and

late

ral

pre

fro

nta

lan

dp

ost

erio

rco

rtic

esJe

ffri

eset

al.

(200

3)P

ET

stu

dy

inve

stig

atin

gb

rain

net

wo

rks

acti

vate

dd

uri

ng

sin

gin

gan

dsp

eaki

ng

20N

on

mu

sici

ans

Sin

gin

gN

ot

spec

ifie

dd

lPF

C(1

)L

eft

hem

isp

her

em

ore

acti

vate

dw

hen

pro

du

cin

gsp

eech

and

righ

tw

hen

pro

du

cin

gm

usi

c;(2

)Si

ngi

ng

wo

rds

did

no

tac

tiva

te‘‘m

irro

r-im

age’

’st

ruct

ure

sin

righ

th

emis

ph

ere,

sugg

esti

ng

mu

ltip

len

eura

ln

etw

ork

sm

ayb

ein

volv

edin

dif

fere

nt

asp

ects

of

sin

gin

g

TA

BL

E2

Con

tin

ued


TA

BL

E2

Con

tin

ued

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Jerd

eet

al.

(201

1)P

ET

stu

dy

exp

lori

ng

stru

ctu

res

un

der

lyin

gw

ork

ing

mem

ory

for

rhyt

hm

and

mel

od

y

10N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)R

hyt

hm

,M

elo

dy

AC

CR

hyt

hm

and

mel

od

yse

emto

hav

eu

niq

ue

neu

ral

pro

cess

ing

net

-w

ork

sin

earl

yst

ages

of

pro

-ce

ssin

gan

din

hig

her

cogn

itiv

ep

roce

ssin

go

fw

ork

ing

mem

ory

Kle

ber

etal

.(2

007)

fMR

Ist

ud

yex

plo

rin

gn

eura

ln

etw

ork

sin

volv

edin

sin

gin

gan

dim

agin

edsi

ngi

ng

16M

usi

cian

sSi

ngi

ng

No

tsp

ecif

ied

AC

C,

vlP

FC

,A

myg

dal

a

(1)

Bro

adra

nge

ofac

tivat

ion

inxc

par

tlyov

erla

pp

ing

cort

ical

and

subc

ortic

alar

eas

wh

enov

ertly

and

imag

inin

gsi

ngi

ng;

(2)

Imag

ined

sin

gin

gac

tivat

edar

eas

invo

lved

inw

orki

ng

mem

ory;

(3)

Em

otio

np

roce

ssin

gar

eas

show

eden

han

ced

activ

atio

nd

uri

ng

imag

ined

sin

gin

gK

leb

eret

al.

(201

0)fM

RI

stu

dy

exp

lori

ng

effe

cto

fvo

cal

mo

tor

skil

lstr

ain

ing

on

fun

ctio

nal

som

ato

sen

sory

acti

vati

on

49B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Sin

gin

gN

ot

spec

ifie

dd

lPF

C(1

)M

usi

ctr

ain

ing

asso

ciat

edw

ith

incr

ease

dac

tiva

tio

nin

dif

fuse

mo

tor,

sen

sory

,fr

on

tal,

par

ieta

l,su

bco

rtic

al,

and

cere

bel

lar

stru

ctu

res;

(2)

Vo

cal

skil

lstr

ain

ing

corr

elat

edw

ith

incr

ease

dac

tivi

tyin

kin

esth

etic

mo

tor

con

tro

l,se

nso

rim

oto

rgu

idan

ce,

and

imp

lici

tm

oto

rm

emo

ryar

eas

Vol. 50, No. 3, Fall 2013 217

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Kn

osc

he

etal

.(2

005)

EE

Gan

dM

EG

stu

dy

exp

lori

ng

neu

ral-

bas

edp

erce

pti

on

of

mu

sica

lp

hra

sest

ruct

ure

12M

usi

cian

sM

usi

cli

sten

ing

(rec

ord

ed)

Mel

od

yA

CC

,O

FCT

imin

gan

dto

po

grap

hy

for

pro

cess

ing

mu

sica

lp

hra

ses

sim

ilar

toth

ose

use

dto

pro

cess

pro

sod

icp

hra

seb

ou

nd

arie

sin

spee

chK

oel

sch

etal

.(2

006)

fMR

Ist

ud

yex

plo

rin

gn

eura

lm

ech

anis

ms

invo

lved

inp

roce

ssin

gp

leas

ant

v.u

np

leas

ant

mu

sic

11N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)N

ot

spec

ifie

dA

myg

dal

a(1

)D

iffe

ren

tst

ruct

ure

san

dn

etw

ork

sac

tiva

ted

wh

enp

roce

ssin

gu

np

leas

ant

v.p

leas

ant

mu

sic;

(2)

Act

ivat

ion

sin

crea

sed

ove

rti

me

du

rin

gp

rese

nta

tio

no

fm

usi

c,in

dic

atin

gti

me

effe

cto

fem

oti

on

pro

cess

ing

Ko

elsc

het

al.

(200

8)fM

RI

stu

dy

inve

stig

atin

gn

eura

ln

etw

ork

sin

volv

edin

per

ceiv

edem

oti

on

alva

len

cech

ord

s

20B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

yA

myg

dal

a(1

)M

usi

c-sy

nta

ctic

aler

rors

acti

vate

dst

ruct

ure

sre

late

dto

emo

tio

nal

pro

cess

ing,

incl

ud

ing

amyg

dal

a;(2

)Ir

regu

lar

cho

rds

wer

eju

dge

dm

ore

un

ple

asan

tco

mp

ared

to‘‘r

egu

lar’

’ch

ord

end

ing

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Lee

etal

.(2

011)

fMR

Ist

ud

yex

plo

rin

gb

rain

regi

on

sim

pli

cati

on

ind

iscr

imin

atio

nm

elo

dic

con

tou

r

12N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)M

elo

dy

AC

C(1

)R

epo

rted

on

3d

isti

nct

cort

ical

area

s,in

clu

din

gA

CC

,w

ho

seac

tivi

tyse

emed

tod

iscr

imin

ate

dif

fere

nt

con

tou

rs;(

2)D

esce

nd

ing

and

asce

nd

ing

par

tso

fth

eco

nto

ur

alte

rb

rain

acti

vati

on

pat

tern

sL

ern

eret

al.

(200

9)fM

RI

stu

dy

exp

lori

ng

dif

fere

nce

sin

neu

tral

net

wo

rks

acti

vate

dw

hen

list

enin

gto

mu

sic

wit

hey

eso

pen

v.cl

ose

d

15N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)N

ot

spec

ifie

dA

myg

dal

a(1

)G

reat

erac

tiva

tio

no

fam

ygd

ala

wh

enli

sten

ing

toem

oti

on

alm

usi

cw

ith

eyes

clo

sed

and

wh

enli

sten

ing

ton

egat

ive

mu

sic;

(2)

Fin

din

gssu

pp

ort

syst

em-b

ased

mo

del

of

per

ceiv

edem

oti

on

alit

yw

ith

amyg

dal

ah

avin

gce

ntr

alro

lein

med

iati

ng

effe

cts

of

con

text

-bas

edp

roce

ssin

gb

yre

cru

itin

g‘‘l

ow

’’(e

.g.,

visc

eral

)an

d‘‘h

igh

’’(e

.g.,

cogn

itiv

e)n

eura

lo

per

atio

ns

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 219

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Lim

b&

Bra

un

(200

8)fM

RI

stu

dy

exp

lori

ng

neu

ral

net

wo

rks

invo

lved

inm

usi

cim

pro

visa

tio

n

6M

usi

cian

sIm

pro

visa

tio

n,

inst

rum

ent

Pla

yN

ot

spec

ifie

dO

FC

,d

lPF

C,

Am

ygd

ala

(1)

Spo

nta

neo

us

imp

rovi

sati

on

char

acte

rize

db

yw

ides

pre

add

eact

ivat

ion

of

lPF

Can

dfo

cal

acti

vati

on

of

mP

FC

,re

gard

less

of

com

ple

xity

,in

dic

atin

gco

gnit

ive

dis

soci

atio

ns

inth

ecr

eati

vep

roce

ss;

(2)

Imp

rovi

sati

on

cau

sed

acti

vati

on

of

cort

ical

sen

sori

mo

tor

area

san

dd

eact

ivat

ion

of

lim

bic

stru

ctu

res

Men

on

&L

evit

in(2

005)

fMR

Ist

ud

yex

plo

rin

gn

eura

ln

etw

ork

sac

tiva

ted

wh

enh

avin

gan

emo

tio

nal

reac

tio

nto

mu

sic

13N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)N

ot

spec

ifie

dO

FC

,A

CC

(1)

Lis

ten

ing

tom

usi

cst

ron

gly

mo

du

late

sac

tivi

tyin

mes

oli

mb

icre

war

d-

pro

cess

ing

net

wo

rk,

incl

ud

ing

the

AC

C,

and

stru

ctu

res

invo

lved

inre

gula

tin

gau

ton

om

ican

dp

hys

iolo

gica

lre

spo

nse

sto

emo

tio

nal

stim

uli

;(2

)M

usi

cli

sten

ing

may

con

nec

taf

fect

ive

and

auto

no

mic

pro

cess

ing

syst

ems

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Mit

ters

-ch

ifft

hal

eret

al.

(200

7)

fMR

Ist

ud

yex

plo

rin

gn

eura

ld

iffe

ren

ces

wh

enli

sten

ing

toh

app

yv.

sad

mu

sic

16N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

AC

C,

Am

ygd

ala

(1)

Dif

fere

nt

neu

ral

net

wo

rks

acti

vate

dw

hen

list

enin

gto

hap

py/

sad

/n

eutr

alm

usi

c;(2

)M

usi

c-el

icit

edem

oti

on

pro

cess

ing

may

inte

grat

eve

ntr

alan

dd

ors

alst

riat

um

(fo

rre

war

dex

per

ien

cean

dm

ove

men

t),

AC

C(f

or

atte

nti

on

),an

dm

edia

lte

mp

ora

llo

bes

(fo

rem

oti

on

app

rais

alan

dp

roce

ssin

g);(

3)R

epo

rted

ano

rder

effe

ctin

acti

vati

on

pat

tern

sw

hen

list

enin

gto

hap

py

mu

sic

firs

tv.

sad

mu

sic

Miz

un

o&

Sugi

shit

a(2

007)

fMR

Ist

ud

yin

vest

igat

ing

neu

ral

corr

elat

esto

pro

cess

ing

mo

de-

ind

uce

dem

oti

on

alre

spo

nse

s

18M

usi

cian

sM

usi

cli

sten

ing

(rec

ord

ed)

Har

mo

ny

AC

C(1

)C

erta

inst

ruct

ure

san

dn

etw

ork

sw

ere

acti

vate

dw

hen

list

enin

gto

tria

ds

wit

ha

ton

alst

ruct

ure

than

tho

sew

ith

ou

t;(2

)A

CC

acti

vate

dd

uri

ng

min

or

and

maj

or

mo

de

con

dit

ion

s

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 221

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Mu

tsh

ule

ret

al.

(201

0)fM

RI

stu

dy

exp

lori

ng

chan

ges

inb

rain

acti

vity

wh

enh

abit

uat

edto

anaf

fect

ive

mu

sica

lst

imu

li

19N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

y,te

mp

ovl

PF

C,

Am

ygd

ala

(1)

Rep

ort

edam

ygd

ala-

cort

ical

net

wo

rks

imp

lica

ted

inh

abit

uat

ion

effe

cts

of

emo

tio

nal

exp

erie

nce

s;(2

)D

iffe

ren

tti

me

scal

eso

fh

abit

uat

ion

coex

ist

wit

hp

erce

pti

on

of

mu

sic

Nak

amu

raet

al.

(199

9)P

ET

and

EE

Gst

ud

yex

plo

rin

gn

eura

ln

etw

ork

sac

tiva

ted

wh

enli

sten

ing

tom

usi

c

8N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

AC

CP

assi

vem

usi

cli

sten

ing

cau

sed

incr

ease

acti

vati

on

inp

ost

erio

rtw

o-t

hir

ds

of

scal

p,

sugg

esti

ng

anin

tera

ctio

nb

etw

een

mu

sic

pro

cess

ing

and

cogn

itiv

ep

roce

sses

Oh

nis

hi

etal

.(2

001)

fMR

Ist

ud

yex

plo

rin

gd

iffe

ren

ces

inac

tiva

tio

np

atte

rns

inm

usi

cian

sv.

no

nm

usi

cian

sd

uri

ng

mu

sic

per

cep

tio

n

28B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Mu

sic

list

enin

g(r

eco

rded

)N

ot

spec

ifie

dd

lPF

CR

epo

rted

left

hem

isp

her

ed

om

inan

cew

hen

list

enin

gto

mu

sic

for

mu

sici

ans,

righ

th

emis

ph

ere

do

min

ance

for

no

nm

usi

cian

s,an

dsi

gnif

ican

td

iffe

ren

cein

deg

ree

of

acti

vati

on

ince

rtai

nar

eas

inm

usi

cian

s,in

clu

din

gle

ftd

lPF

C

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Pal

lese

net

al.

(200

9)fM

RI

stu

dy

inve

stig

atin

gch

ange

sin

bra

inac

tiva

tio

nw

hen

cogn

itiv

ely

and

emo

tio

nal

lyp

roce

ssin

gaf

fect

ive

stim

uli

10N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

yrt

.A

myg

dal

a(1

)N

ote

dd

iffe

ren

ces

inac

tiva

tio

np

atte

rns

du

rin

gm

usi

cli

sten

ing

wit

han

dw

ith

ou

tw

ork

ing

mem

ory

task

;(2

)In

som

ere

gio

ns,

the

grea

ter

the

wo

rkin

gm

emo

ryta

skth

ela

rger

the

dec

reas

e;(3

)T

ask-

rela

ted

dec

reas

esm

ayb

efu

rth

eraf

fect

edb

yem

oti

on

alim

pac

to

fm

usi

cP

alle

sen

etal

.(2

005)

fMR

Ist

ud

yex

plo

rin

gn

eura

ln

etw

ork

sin

volv

edin

pro

cess

ing

emo

tio

nal

resp

on

ses

toch

ord

sin

mu

sici

ans

and

no

nm

usi

cian

s

21B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

yA

myg

dal

a(1

)N

eura

lp

roce

ssin

gin

emo

tio

n-r

elat

edb

rain

area

sca

nb

eac

tiva

ted

by

sin

gle

cho

rds;

(2)

Em

oti

on

pro

cess

ing

was

enh

ance

din

abse

nce

of

cogn

itiv

ere

qu

irem

ents

;(3

)M

usi

cian

san

dn

on

mu

sici

ans

do

no

td

iffe

rin

neu

ral

pro

cess

ing

of

sin

gle

cho

rds

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 223

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Pal

lese

net

al.

(201

0)fM

RI

stu

dy

exp

lori

ng

dif

fere

nce

sb

etw

een

mu

sici

ans

and

no

nm

usi

cian

sin

wo

rkin

gm

emo

ryo

fm

usi

cal

sou

nd

task

21B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Mu

sic

list

enin

g(r

eco

rded

)H

arm

on

yA

CC

,L

PF

C(1

)M

usi

cian

sp

erfo

rmed

bet

ter

on

cogn

itiv

eta

sks;

(2)

Mu

sici

ans

had

incr

ease

dac

tiva

tio

nin

area

sim

plic

ated

inat

ten

tio

nan

dco

gnit

ive

con

tro

l,es

pec

ially

inri

ght

hem

isp

her

e;(3

)R

elat

ion

-sh

ipb

etw

een

task

per

form

ance

and

mag

nit

ud

eo

fre

spo

nse

mo

rep

osi

tive

inm

usi

cian

sP

erry

etal

.(1

999)

PE

Tsc

anin

vest

igat

ing

neu

ral

acti

vati

on

pat

tern

sas

soci

ated

wit

hsi

ngi

ng

13N

on

mu

sici

ans

Mu

sic

list

enin

g(r

eco

rded

),Si

ngi

ng

No

tsp

ecif

ied

AC

C(1

)R

epo

rted

on

aco

mp

lex,

dis

trib

ute

dn

etw

ork

of

cort

ical

acti

vati

on

pat

tern

sd

uri

ng

sin

gin

gp

rod

uct

ion

;(2)

Sim

ple

,ch

ant-l

ike

sin

gin

gac

tiva

tes

sim

ilar

net

wo

rks

assp

eech

wit

hso

me

hem

isp

her

icd

iffe

ren

ces

inm

oto

ran

dau

dit

ory

regi

on

sR

uiz

etal

.(2

009)

EE

Gst

ud

yex

plo

rin

gn

eura

lco

rrel

ates

asso

ciat

edw

ith

exec

uti

veco

ntr

ol

du

rin

gp

ian

op

layi

ng

19M

usi

cian

sIn

stru

men

tP

lay

No

tsp

ecif

ied

AC

C(1

)E

rro

rm

on

ito

rin

gn

etw

ork

s,ge

ner

ated

by

AC

C,

pro

cess

eser

rors

70m

sp

rio

rto

them

;(2

)R

epo

rted

on

dif

fere

nt

con

trib

u-

tio

ns

of

aud

ito

ryan

dso

mat

o-

sen

sory

info

rmat

ion

toer

ror

mo

nit

ori

ng;

(3)

Au

dit

ory

info

rmat

ion

mo

du

late

der

ror

pro

cess

ing

po

st-e

xecu

tio

n

TA

BL

E2

Con

tin

ued


Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Sato

het

al.

(200

1)P

ET

scan

inve

stig

atin

gd

iffe

ren

ces

mu

sica

lm

od

eh

aso

nn

eura

lac

tiva

tio

np

atte

rns

9M

usi

cian

sM

usi

cli

sten

ing

(rec

ord

ed)

Har

mo

ny

OF

C(1

)A

tten

din

gto

mel

od

iclin

ere

-cr

uit

edar

eas

rela

ted

tose

lect

ive

atte

nti

on

and

ton

al-v

erb

alas

soci

atio

ns

(e.g

.,O

FC);

(2)

Att

end

ing

toh

arm

on

yac

tiva

ted

regi

on

sre

late

dto

emo

tio

nal

pro

cess

ing

and

mem

ory

Sud

aet

al.

(200

8)O

Tsy

stem

stu

dy

exp

lori

ng

effe

cto

fM

oza

rt’s

mu

sic

on

spat

ial-r

easo

nin

gab

ilit

y

10N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

No

tsp

ecif

ied

dlP

FC

Exp

osu

reto

Moz

art’

sm

usi

cen

han

ced

per

form

ance

onin

telli

gen

cete

sts

and

reve

aled

diff

eren

tac

tivat

ion

pat

tern

sin

area

sim

plic

ated

insp

atia

l-te

mp

oral

reas

onin

g(e

.g.,

dlP

FC)

Th

aut

etal

.(2

009)

PE

Tsc

anex

plo

rin

gac

tiva

tio

np

atte

rns

asso

ciat

edw

ith

rhyt

hm

icau

dit

ory

mo

tor

syn

chro

niz

atio

n

9N

ot

spec

ifie

dM

usi

cli

sten

ing

(rec

ord

ed)

Rh

yth

md

lPF

C(1

)R

epo

rted

on

stru

ctu

res

invo

lved

ind

iffe

ren

tas

pec

tso

fm

oto

rsy

nch

ron

izat

ion

;(2

)F

ind

ings

sugg

est

dis

tin

ct,

fun

ctio

nal

cort

ico

-cer

ebel

lar

circ

uit

sse

rve

dif

fere

nt

asp

ects

of

rhyt

hm

icsy

nch

ron

izat

ion

,co

nsc

iou

san

dsu

bco

nsc

iou

sre

spo

nse

tote

mp

ora

lst

ruct

ure

,an

dco

nsc

iou

sm

on

ito

rin

go

frh

yth

mic

pat

tern

trac

kin

g

TA

BL

E2

Con

tin

ued

Vol. 50, No. 3, Fall 2013 225

Au

tho

rD

escr

ipti

on

of

stu

dy

Stu

dy

char

acte

rist

ics

Ou

tco

mes

NP

arti

cip

ant

mu

sica

lab

ilit

yT

ype

of

exp

erie

nce

(s)

Mu

sic

char

acte

rist

icN

eura

lst

ruct

ure

(s)

Gen

eral

fin

din

gs

Vo

gtet

al.

(200

7)fM

RI

stu

dy

exp

lori

ng

effe

cto

fp

ract

icin

go

nn

eura

lac

tiva

tio

np

atte

rns

32B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Inst

rum

ent

Pla

yN

ot

spec

ifie

dd

lPF

C(1

)M

irro

rn

euro

nsy

stem

mo

rest

ron

gly

acti

vate

dd

uri

ng

ob

ser-

vati

on

of

no

n-p

ract

iced

item

s;(2

)L

eft

dlP

FCse

lect

ivel

yin

volv

edd

uri

ng

ob

serv

atio

nan

dm

oto

rp

rep

of

no

n-

pra

ctic

edch

ord

sZ

arat

e&

Zat

orr

e(2

008)

fMR

Ist

ud

yex

plo

rin

gd

iffe

ren

ces

inau

dio

-vo

cal

inte

grat

ion

inm

usi

cian

sv.

no

nm

usi

cian

s

24B

oth

mu

sici

ans

and

no

nm

usi

cian

s

Sin

gin

gP

itch

AC

C(1

)Si

nge

rsm

ore

accu

rate

than

no

n-s

inge

rsin

sin

gin

g(w

ith

sam

en

eura

ln

etw

ork

sre

cru

ited

),at

ign

ori

ng

shif

tin

gfe

edb

ack

(wit

hd

iffe

ren

tn

eura

ln

etw

ork

s),

and

sam

eat

com

pen

sate

task

(wit

hd

iffe

ren

tn

eura

ln

etw

ork

s);

(2)

Au

tho

rsp

rop

ose

two

neu

ral

sub

stra

tes

for

aud

io-v

oca

lin

tegr

atio

nZ

arat

eet

al.

(201

0)fM

RI

stu

dy

inve

stig

atin

gn

eura

ln

etw

ork

sin

volv

edin

volu

nta

ryv.

invo

lun

tary

pit

chre

gula

tio

n

9M

usi

cian

sSi

ngi

ng

Pit

chA

CC

(1)

Sin

gers

less

able

toig

no

rem

ino

rp

itch

-sh

ift

dif

fere

nce

sth

anm

ore

no

tice

able

on

es;(

2)C

om

pen

sate

task

recr

uit

edfu

nct

ion

ally

-co

nn

ecte

dn

eura

ln

etw

ork

,in

clu

din

gA

CC

;(3

)L

arge

rvo

calc

orr

ecti

on

sap

pea

rto

be

un

der

cort

ical

con

tro

l

TA

BL

E2

Con

tin

ued


TA

BL

E3

Sum

mar

yof

Mai

nFi

ndi

ngs

and

Cli

nic

alC

onsi

dera

tion

s

Neu

ral

stru

ctu

reSu

mm

ary

of

fin

din

gsC

lin

ical

con

sid

erat

ion

s

Am

ygd

ala

Act

ivat

ion

:N

occ

urr

edw

hen

list

enin

gto

min

or/

dis

son

ant/

neg

ativ

e/u

np

leas

ant

mu

sic,

du

rin

gan

un

exp

ecte

dev

ent,

du

rin

gm

usi

cli

sten

ing

inge

ner

alN

cou

ldb

em

od

ula

ted

by

asi

ngl

ech

ord

chan

geN

incr

ease

dw

hen

list

enin

gto

mu

sic

wit

hey

escl

ose

d,

wh

enp

airi

ng

mu

sic

and

visu

alst

imu

liD

eact

ivat

ion

:N

occ

urr

edd

uri

ng

mu

sic

imp

rovi

sati

on

,w

hen

list

enin

gto

ple

asan

tm

usi

cR

igh

tam

ygd

ala:

Nwas

recr

uit

edd

uri

ng

‘‘sad

’’m

usi

c,im

plic

ated

inea

rly

resp

on

ses

toch

ord

vio

lati

on

s,ex

hib

its

cho

rd-d

epen

den

tre

spo

nse

sN

was

acti

vate

dm

ore

stro

ngl

yd

uri

ng

aud

iovi

sual

con

dit

ion

Nac

tivi

tyin

crea

sed

wit

hex

po

sure

ove

rti

me

Lef

tam

ygd

ala:

Nac

tivi

tyin

crea

sed

wh

enlis

ten

ing

tom

usi

cw

ith

ah

igh

ern

egat

ive

vale

nce

Lo

ng-

term

hab

itu

atio

nef

fect

was

no

ted

To

faci

lita

teE

R:

Nlis

ten

tom

usi

cth

ecl

ien

tco

nsi

der

sp

leas

ant

or

hap

py

Nin

corp

ora

tem

usi

cim

pro

visa

tio

nN

refr

ain

fro

mh

avin

gsu

dd

enan

du

nex

pec

ted

mu

sica

lev

ents

(e.g

.,ab

rup

tch

ord

chan

ges,

sud

den

dyn

amic

chan

ges,

etc.

,)N

con

sid

erth

eef

fect

of

pai

rin

gm

usi

cw

ith

avi

sual

stim

ulu

sN

do

no

tli

sten

tom

usi

cw

ith

eyes

clo

sed

An

teri

or

Cin

gula

teC

ort

ex(A

CC

)A

ctiv

atio

n:

No

ccu

rred

du

rin

gvo

lun

tary

pit

chco

rrec

tio

nan

dm

usi

cd

iscr

imin

atio

nta

sks

No

ccu

rred

wh

enli

sten

ing

toch

ord

vio

lati

on

s,w

hen

list

enin

gto

fam

ilia

r/fa

vora

ble

/en

erge

tic

mu

sic,

du

rin

go

vert

and

imag

ined

sin

gin

gNo

ccu

rred

wh

enlis

ten

ing

tod

isso

nan

tch

ord

s(m

usi

cian

so

nly

)N

incr

ease

dd

uri

ng

sin

gin

gan

dm

usi

cli

sten

ing

task

sNw

asm

ore

pro

no

un

ced

inm

usi

cian

sco

mp

ared

ton

on

mu

sici

ans

Nin

crea

sed

inth

ose

face

dw

ith

am

ism

atch

bet

wee

nin

div

idu

alan

dgr

ou

po

pin

ion

,is

corr

elat

edw

ith

son

gli

kab

ilit

y

To

faci

lita

teE

R:

Nli

sten

tom

usi

ccl

ien

tco

nsi

der

sfa

mil

iar

and

pre

ferr

edN

enga

gecl

ien

tin

acti

vem

usi

cm

akin

g,su

chas

sin

gin

go

rm

usi

cim

pro

visa

tio

nNe

nga

gecl

ien

tin

atte

nd

ing

tosp

ecif

icm

usi

calc

ues

,su

chas

ph

rase

so

rm

elo

dic

lin

es

Vol. 50, No. 3, Fall 2013 227

Neu

ral

stru

ctu

reSu

mm

ary

of

fin

din

gsC

lin

ical

con

sid

erat

ion

s

Nm

ayb

em

od

e-d

epen

den

t,i.e

.,p

atte

rns

chan

geb

ased

on

mo

des

(res

ult

sm

ixed

)N

sho

wed

dif

fere

nt

pat

tern

sw

hen

per

ceiv

ing

asce

nd

ing

and

des

cen

din

gm

elo

dic

con

tou

rsth

anw

hen

pro

cess

ing

mu

sica

lp

hra

seb

ou

nd

arie

sN

incr

ease

din

ven

tral

AC

Cve

rsu

sd

ors

alA

CC

du

rin

gm

usi

cli

sten

ing

Rig

ht

AC

C:

Nw

asac

tiva

ted

du

rin

gm

usi

cim

pro

visa

tio

nta

skL

eft

AC

CN

was

acti

vate

dd

uri

ng

rhyt

hm

-an

dm

elo

dy-

bas

edw

ork

ing

mem

ory

task

s,w

hen

list

enin

gto

favo

rab

le/

ener

geti

cm

usi

cN

was

acti

vate

dm

ore

stro

ngl

yfo

rrh

yth

mO

rbit

ofr

on

tal

Co

rtex

(OF

C)

Act

ivat

ion

:N

occ

urr

edw

hen

atte

nd

ing

toa

sin

gle

mu

sica

lco

mp

on

ent

(e.g

.,m

elo

dy)

,d

etec

tin

gp

hra

seb

ou

nd

arie

s,p

roce

ssin

gty

pes

of

cho

rds,

resp

on

din

gto

syn

tact

ical

irre

gula

riti

esN

occ

urr

edw

hen

mu

sic

list

enin

gre

gard

less

of

pre

fere

nce

/va

len

ce(r

esu

lts

mix

ed)

Nw

asst

ron

ger

du

rin

gm

usi

cli

sten

ing

than

spee

chli

sten

ing

Nre

late

dto

son

gli

kab

ilit

y,w

ith

incr

ease

dac

tiva

tio

nfo

rn

on

-like

dso

ngs

Rig

ht

OF

C:

Nac

tiva

tio

nn

egat

ivel

yco

rrel

ated

wit

hch

ord

inst

abil

ity

Nac

tiva

tio

nin

crea

sed

com

par

edto

the

left

OF

Cw

hen

pro

cess

ing

emo

tio

nal

ly-s

alie

nt

mu

sic

To

faci

lita

teE

R:

Nli

sten

tom

usi

ccl

ien

tco

nsi

der

sfa

mil

iar

and

pre

ferr

edNe

nga

gecl

ien

tin

atte

nd

ing

tosp

ecif

icm

usi

cal

cues

,su

chas

ph

rase

so

rm

elo

dic

lin

esN

cho

ose

mu

sic

that

use

sst

able

cho

rds,

i.e.

,re

frai

nfr

om

too

mu

chd

isso

nan

ce

TA

BL

E3

Con

tin

ued


Neu

ral

stru

ctu

reSu

mm

ary

of

fin

din

gsC

lin

ical

con

sid

erat

ion

s

Lat

eral

Pre

fro

nta

lC

ort

ex(l

PF

C),

incl

ud

ing

Do

rso

late

ral

Pre

fro

nta

lC

ort

ex(d

lPF

C)

and

Ven

tro

late

ral

Pre

fro

nta

lC

ort

ex(v

lPF

C)

lPF

Cac

tiva

tio

n:

Nw

asst

ron

ger

du

rin

ga

mu

sic-

bas

edw

ork

ing

mem

ory

task

and

inre

spo

nse

ton

egat

ive

mu

sic

pai

red

wit

hvi

sual

stim

uli

Nco

rrel

ated

wit

hso

ng

lika

bil

ity

dlP

FC

acti

vati

on

:N

occ

urr

edw

hen

ob

serv

ing

and

pre

par

ing

top

lay

no

n-p

ract

iced

cho

rds,

wh

enli

sten

ing

toM

oza

rtm

usi

c,w

hen

list

enin

gto

mu

sic

wit

hey

eso

pen

bu

tn

ot

clo

sed

Nin

crea

sed

wh

ensy

nch

ron

izin

gto

tem

po

chan

ges

du

rin

ga

mo

tor-

rhyt

hm

syn

chro

niz

atio

nta

skN

pat

tern

sm

ixed

for

mu

sic

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Levitin, 2005; Nakamura et al., 1999) and it was more pronouncedin musicians compared to nonmusicians (Pallesen et al., 2010;Zarate & Zatorre, 2008). ACC activation was correlated withoverall song likability (Berns & Moore, 2012) and increased inindividuals who disliked a song, yet were sensitive to the song’spopularity; the researchers attributed this ACC activation to amismatch between individual and societal opinion (Berns, Capra,Moore, & Noussair, 2010). Reports of activation patterns based onmusical modes are mixed, with studies reporting increasedactivation when listening to happy music or music that elicitschills (Blood & Zatorre, 2001; Mitterschiffthaler et al., 2007),when listening to music in a minor mode as compared to a majormode (Green et al., 2008), or when listening to music in either amajor or minor mode (Mizuno & Sugishita, 2007). Differentactivation patterns were reported when listeners perceivedascending and descending melodic contours (Lee, Janata, Frost,Hanke, & Granger, 2011) and when processing musical phraseboundaries (Knosche et al., 2005). In addition, the ACC wasactivated during both overt and imagined singing (Kleber,Birbaumer, Veit, Trevorrow, & Lotze, 2007) and during singingimprovisation (Brown, Martinez, & Parsons, 2006). The left ACCwas activated during rhythm- and melody-based working memorytasks, though more strongly for rhythm (Jerde, Childs, Handy,Nagode, & Pardo, 2011) and when listening to favorable orenergetic music (Brown, Martinez, & Parsons, 2004), whereas theright ACC was activated during a free generation task (deManzano & Ullen, 2012). Finally, there was increased activity inthe ventral ACC when listening to music compared to the dorsalACC (Green et al., 2008). To summarize, ACC activation occurredwhen listening to preferred music, when engaged in an activemusic-making experience, and when attending to specificcharacteristics of the music stimulus.

Orbitofrontal cortex (OFC). The OFC was activated whenlistening to preferred music (Menon & Levitin, 2005) and in aseparate study was more strongly activated when listening to musicas compared with listening to speech (Callan et al., 2006). OFCactivation related to emotional valence were mixed, with studiesreporting that listening to unpleasant or negative music activatedthe OFC (Flores-Gutierrez et al., 2007) as well as listening topleasant music (Blood & Zatorre, 2001). Several studies reported


activation patterns related to attending to musical characteristics.The OFC was activated when attending to a single musicalcomponent, such as a melodic line (Satoh, Takeda, Nagata,Hatazawa, & Kuzuhara, 2001), when detecting phrase boundaries(Knosche et al., 2005), when processing types of chords (Fujisawa& Cook, 2011), and when responding to syntactical irregularities(James et al., 2008). OFC activation is reported to be related tosong likability (Berns et al., 2010; Berns & Moore, 2012), withincreased activation for nonliked songs (Berns & Moore, 2012).The right OFC may be implicated as a link between affective andcognitive neural systems that are engaged during music listening(Menon & Levitin, 2005). Its activation is negatively correlatedwith chord instability (e.g., dissonant chords) (Fujisawa & Cook,2011) and it showed increased activation compared to the leftOFC when processing emotionally-salient music (e.g., eitherunpleasant/negative music or pleasant music) (Blood & Zatorre,2001; Flores-Gutierrez et al., 2007). To summarize, OFC activationoccurred when listening to preferred music, when listening tofamiliar music, and when attending to specific characteristics ofthe music stimulus.

Lateral prefrontal cortex (PFC). The majority of studies thatreported on prefrontal cortex activity focused on either thedorsolateral prefrontal cortex (dlPFC) or the ventrolateralprefrontal cortex (vlPFC). Those that reported on the lateralPFC noted enhanced activity during a music-based workingmemory task (Pallesen et al., 2010), stronger activation inresponse to negative music paired with visual stimuli, but notwithout visual stimuli (Eldar et al., 2007), and activation that wascorrelated with song likability (Berns et al., 2010).

Dorsolateral prefrontal cortex (dlPFC). The dlPFC was activat-ed during the observation and motor preparation of nonpracticedchords (Vogt et al., 2007), when listening to Mozart music (Suda,Morimoto, Obata, Koizumi, & Maki, 2008), and when listening tomusic with eyes open, but not closed (Lerner et al., 2009).Increased activation was noted when synchronizing to tempochanges during a motor-rhythm synchronization task (Thaut etal., 2009) and mixed results were reported when exploring dlPFCactivation during a music improvisation task, which either causedactivation of the dlPFC (Bengtsson, Csıkszentmihalyi, & Ullen,2007) or deactivation (Limb & Braun, 2008). The impact of music

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training on dlPFC activation also had mixed results, with onestudy noting that musicians had stronger dlPFC activation (Ohnisiet al., 2001) and another reporting that college-aged music majorshad stronger dlPFC activation than laypersons, but not profes-sional musicians (Kleber, Veit, Birbaumer, Gruzelier, & Lotze,2010). The right dlPFC was activated during singing (Jeffries,Fritz, & Braun, 2003), and during music discrimination tasks(Brown & Martinez, 2007; Hugdahl et al., 1999). The left dlPFCwas activated during a free generation task, i.e., a musicimprovisation task (de Manzano & Ullen, 2012).

Ventrolateral prefrontal cortex (vlPFC). The vlPFC was activat-ed when listening to musical triads (Mutshuler et al., 2010) and itmay be implicated in processing syntactical violations in music(Janata, 2009). In addition, vlPFC activation occurred during animagined singing task, but not when overtly singing, perhapsreflecting its role in emotional recall (Kleber et al., 2007).

To summarize, lateral PFC activation occurred when listeningto preferred or familiar music, when engaged in an active music-making experience, and when attending to specific characteristicsof the music stimulus.

Discussion

The purpose of this exploratory review was to examine theeffects of music on neural structures implicated in emotionregulation and to create preliminary clinical considerations basedon this synthesis. Although the impact of music on emotionprocessing has long been of interest, this is the first attempt tosystematically review and synthesize research specifically investi-gating the neural effect of music on emotion regulation. Resultsindicated that there are certain musical characteristics andexperiences that produce desired neural activation patternsimplicated in emotion regulation. From a clinical perspective,understanding this link between musical elements and the targetgoal—in this case, emotion regulation—helps the music therapistmake informed decisions about the Therapeutic Function ofMusic (TFM). Hanson-Abromeit (2013) defines the TFM as ‘‘thedirect relationship between the treatment goal and the explicitcharacteristics of the musical elements, informed by a theoreticalframework and/or philosophical paradigm in the context of aclient’’ (p. 130). In other words, having an explicit understanding


of why and how music affects a desired change informs theintentional, therapeutic use of music in clinical practice. For thepurposes of this study, it involves having an explicit understandingof what musical characteristics and experiences impact a person’sability to regulate his or her emotions and how.

Emotion regulation is characterized by increased activation inthe cognitive control and monitoring areas—the anterior cingulatecortex (ACC), orbitofrontal cortex (OFC), and lateral prefrontalcortex (PFC)—which leads to decreased activation of the amygdala(Gyurak et al., 2011; Ochsner & Gross, 2005; McRae et al., 2010;Rempel-Clower, 2007). Overall, results indicated that there arecertain music characteristics and music experiences that producesuch activation patterns. For example, listening to music consideredpleasant or happy activated the ACC (Blood & Zatorre, 2001; Brownet al., 2004; Mitterschiffthaler et al., 2007), the OFC (Berns & Moore,2012; Berns et al., 2010; Berns & Moore, 2012; Blood & Zatorre,2001; Flores-Gutierrez et al., 2007), and decreased activation in theamygdala (Blood & Zatorre, 2001; Koelsch et al., 2006). Similaractivation patterns were found when listening to music, regardless ofemotional meaning for the listener (Callan et al., 2006; Menon &Levitin, 2005; Mizuno & Sugishita, 2007; Nakamura et al., 1999) andwhen singing (Kleber et al., 2007; Perry et al., 1999). These patternswere also reported when musically-trained individuals were impro-vising (Bengtsson et al., 2007; Brown et al., 2006; Limb & Braun,2008). It should be noted, though, that one study reporteddeactivation in the dlPFC during improvisation, which the authorsattributed to its role in providing a framework for goal-directedbehaviors, a role not needed when improvising (Limb & Braun,2008). Although it is premature to make direct clinically-basedgeneralizations until future research is conducted, these resultsprovide preliminary evidence supporting the use of music listening,singing, and improvisation to facilitate emotion regulation.

In addition to desired neural activation patterns for emotionregulation, there can also be undesired activation patterns.Results of this review indicated certain musical characteristicsand experiences that produced increased activation patterns inthe amygdala, namely listening to music that was minor,dissonant, negative, or unpleasant (Koelsch et al., 2006; Lerneret al., 2009; Mitterschiffthaler et al., 2007; Pallesen et al., 2005)and changing chords (Pallesen et al., 2009), especially in an

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unexpected way, as with chord violations (James et al., 2008;Koelsch et al., 2008). In addition, listening to music with eyes closedincreased the amygdala’s activity (Lerner et al., 2009) as didproviding a more complex sensory stimulus by pairing music with avisual stimulus (Baumgartner et al., 2006; Dyck et al., 2011; Eldar etal., 2007). A primary function of the amygdala is to assessemotionally-salient sensory information. If the sensory informationis determined to be unthreatening, amygdala activity decreases.The findings from this review are congruent with this primaryfunction, as they indicate that the amygdala is processing andassessing music stimuli that is new, (e.g., chord changes),unexpected (e.g., chord violations), or complex (e.g., simultaneousaural and visual input). Thus from a clinical perspective, researchalso indicates characteristics of music and music experiences thatshould be avoided when trying to help a person regulate and shifthis or her physiological and emotional state. Based on this review,these include avoiding music that is minor, dissonant, orconsidered unpleasant, as well as avoiding unexpected musicalevents (e.g., sudden dynamic changes), frequent chord changes,and listening to music with eyes closed. It should be noted that thisprocess implies musically-facilitating a shift to a comfortable state ofarousal, back to homeostasis. There may be times when it isclinically indicated to maintain or increase amygdala activation,thus maintaining or intensifying the emotional experience. Thismay involve the incorporation of music and music experiences thatare considered unpleasant, with frequent chord changes andunexpected musical events. Future research and systematic reviewsare recommended to explore and address this phenomenon.

The roles of the cognitive control and monitoring areas aremore complex than that of the amygdala; therefore, theiractivation patterns associated with music characteristics andexperiences are less straightforward. Results indicated that, forthe most part, all the music experiences produced diffuseactivation in those three areas. However, unlike findings relatedto amygdala functioning, they provided no indication of the typesof music and experiences to use or avoid when facilitatingemotion regulation. One possible explanation for this lack ofclarity is that those structures are implicated in other tasks. Forexample, the ACC is thought to be involved in processinguncertainty and conflict (Brown & Martinez, 2007; Mizuno &


Sugishita, 2007; Pallesen et al., 2010), error monitoring (Ruiz etal., 2009), and response inhibition and selection (de Manzano &Ullen, 2012). Given that music is a complex stimulus, it can behypothesized that once it has been determined that the musicstimulus is not a threat—i.e., the emotion-processing aspect thatresults in decreasing amygdala activity—these neural structuresattend to processing other aspects of the music stimulus.

Another possible explanation relates to their involvement inanother emotion regulation-related process, attention. More specif-ically, this refers to their possible involvement in processing music-influenced changes in attention that facilitate emotion regulation.As indicated previously, a common process involved in emotionregulation is the use of strategies that can create a new emotionalresponse or change a current one (Gyurak et al., 2011; McRae et al.,2010; Ochsner & Gross, 2005). Successful strategies either changehow we interpret the meaning of a situation or alter how we attendto a situation (McRae et al., 2010). All three cognitive control areasexplored in this review are thought to be implicated in networksinvolved in attention, a role reflected in the results of several studiesincluded in this review (de Manzano & Ullen, 2012; Jerde et al.,2011; Knosche et al., 2005; Satoh et al., 2001). One commondenominator in those studies was that the participants wereinstructed to focus on a specific cognitive task related to the musicstimulus, such as processing the melodic phrase structure (Knoscheet al., 2005) or attending to the melodic line and harmonic changes(Satoh et al., 2001). In other words, the participants were instructedto attend to the music stimulus in a different, more analytical way.From a clinical standpoint, one emotion regulation strategy theseresults suggest is to instruct the client to attend to a specificcharacteristic of the music (e.g., the melodic line, a musical cue,etc.), thus removing their focus from the emotional event. Thisstrategy is analogous to using music to reduce pain perception, acommon phenomenon reported in the medical literature (Fra-tianne et al., 2001; Tan, Yowler, Super, & Fratianne, 2010), andsupports the use of effortful, explicit strategies when facilitatingemotion regulation (Gyurak et al., 2011).

Limitations

One major limitation of this review is the lack of interventionreporting in the included studies, specifically that related to the

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music stimulus used. There has been a call in the literature toinclude clear and detailed explanations of the interventions usedand why that intervention was selected (Robb, Burns, &Carpenter, 2011). However, many of the studies included in thisreview were not specific as to the type of music used, the names ofmusical pieces, the structure and characteristics of the musicstimulus, whether the music was original or improvised, or theinstrument(s) that were used. This lack of explicit interventionreporting is problematic for two reasons. One, it makes it difficultto understand the mechanisms underlying the neural activations,thus decreasing the reliability of the results. In other words, itmakes it difficult to explore and understand what characteristicsof the music stimulus are responsible for the desired activationpatterns. Two, without a clear understanding of the musicstimulus used, it is difficult to replicate the stimulus in futurestudies, as well as in clinical practice.

There are other limitations in this review that affect thegeneralizability of the findings. First, as the inclusion criteriastipulated that study participants be typically-developing humans,studies included in this review reflect a nonclinical populationand findings may not easily generalize to clinical populations.Second, the majority of the studies utilized Western music as thestimulus (Table 1), making it unclear how these findings mightgeneralize from a multicultural perspective. As such, the resultsand clinical implications of this review should be consideredpreliminary and should be interpreted with caution. Futureresearch can explore the effect music has on emotion regulationin clinical populations and through a multicultural perspective.Furthermore, clearer intervention reporting is needed in futureresearch to inform the translation of research to clinical practice.

Conclusions

This systematic review indicates that music experiences mayhave an impact on emotion regulation. Furthermore, the resultsprovide preliminary guidelines for music characteristics andspecific strategies that might assist in the emotion regulationprocess when using music as an intervention strategy. Theseguidelines include using music considered happy and pleasant,with predictable, consonant harmonies. In addition, the musictherapy clinician should be aware of multimodal implications,


such as asking a client to close his or her eyes or pairing musicwith a visual stimuli, as these might heighten the emotionalresponse. Music listening, singing, and improvisation may assist infacilitating emotion regulation, as might instructing the client toattend to another task related to the music stimulus, such asfocusing on noticing harmonic changes. As noted, these arepreliminary guidelines and more research is needed. Futureresearch can further explore the Therapeutic Function of Music,teasing apart the different elements of music (e.g., pitch, rhythm,harmony, melody, etc.) and studying their role, if any, on emotionregulation (Hanson-Abromeit, in press). Following that, clinicalstudies are needed to move this exploration from the theoreticalrealm to functional, clinical use. This review provides preliminarysupport for the use of music to facilitate emotion regulation, butstudies are also needed to explore the clinical efficacy of musicinterventions on emotion regulation.

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