Music and Brain Plasticity

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Subject: Psychology,CognitivePsychologyOnlinePublicationDate: Feb2015

DOI: 10.1093/oxfordhb/9780198722946.013.23

MusicandBrainPlasticity SimoneDallaBellaTheOxfordHandbookofMusicPsychology,SecondEdition(Forthcoming)EditedbySusanHallam,IanCross,andMichaelThaut

OxfordHandbooksOnline

AbstractandKeywords

Learningandmakingmusicarecomplexmultimodalactivitieswhichputahighloadonourbrainfunctioning.Theyengageperceptual,cognitive,motor,andsensorimotorprocessesandtheassociatedneuronalcircuitries.Long-andshort-termmusicaltrainingandpracticecanshapebrainstructureandfunctions.Inthischapter,evidencesupportingtheneuroplasticeffectsofmusicisreviewed,byexaminingthedifferencesinmusicians’andnonmusicians’brains,intermsofstructureandfunction.Inaddition,theeffectsofmusicaltrainingandearlymusicexperienceonbraindevelopmentareexamined.Conclusionsaredrawnontheroleplayedbymusic-drivenneuroplasticityforthepurposesofneuroprotectionandrehabilitation.

Keywords:music,brainplasticity,learning,perception,action,sensorimotorprocesses,development,neurorehabilitation

Introduction

Musicaltrainingandperformanceareanidealhumanmodelforexaminingthebraineffectsoflearningspecializedsensorimotorskills.Theyarebothexamplesofmultisensoryexperiencesthatputahighloadonoursensory,motor,andcognitivesystems.Yearsofdedicatedandconstantpracticearenecessarytomastertheskillsdisplayedbyprofessionalmusicians.Playingamusicalinstrumentistypicallyinitiatedearlyduringdevelopment,andproficientperformanceinadultsisgroundedinextensivelearningandpracticeofavarietyofskillsoverthecourseofamusician’slifetime.Acquiringandmaintainingmusicalskillshasimportantandquantifiableeffectsonbrainnetworkingandfunctioning,whicharemanifestationsofbrainplasticity.Incognitiveneurosciences,theterm“brainplasticity”generallyreferstothemalleabilityofthehumanbrain,whichinresponsetoexperienceortrainingpartlymodifiesitsstructureand/orfunctions.Thesechangesarenotconfinedtothedevelopingbrain,but,albeitlesspervasive,theyarealsovisibleinadults(e.g.,WanandSchlaug,2010).Brainplasticityismostimportantwhenthetrainingtaskismeaningfultothesubject(e.g.,Jenkins,Merzenich,Ochs,AllardandGuic-Robles,1990),afactwhichisparticularlyrelevantfortargetedmusicaltraining,andismodulatedbythedegreeofawarenessofsensoryinformationoractionplanning(e.g.,Pascual-Leone,GrafmanandHallett,1994).Moreover,brainplasticityisassociatedwithbehavioraleffectsandisunderpinnedbyprocessesatthecellularandmolecularlevel(e.g.,BuonomanoandMerzenich1998).

Inthischaptertheeffectsofmusicaltrainingandpracticeonbrainstructureandfunctioningarereviewed.Particularattentionispaidtotheeffectsofmusicaltrainingintheadultmusician,andtothemechanismsunderlyingmusiclearningduringdevelopment.

WhereMusicians’andNonmusicians’BrainsDiffer

Afirstapproachtoidentifytheeffectsofmusicaltrainingonbrainplasticityistocompareadultmusiciansand

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nonmusicians.Plasticchangesassociatedwithshort-termandlong-termmusicaltrainingencompassdifferentlevelsofbrainorganizationspanningprimaryperceptualandmotorregionstosensorimotorintegrationandcross-modalassociationareas(forreviews,seeBarrett,Ashley,StraitandKraus,2013;HabibandBesson,2009;HerholzandZatorre,2012;Jäncke,2009;Merrett,PeretzandWilson,2013;Münte,AltenmüllerandJäncke,2002;Rauschecker,2001;StraitandKraus,2014;WanandSchlaug,2010;ZatorreandMcGill,2005).

StructuralandFunctionalDifferencesintheBrainsofMusiciansandNonmusicians

Importantdifferencesbetweenmusiciansandnonmusiciansarevisibleatdifferentlevelsoftheauditorypathway,fromthebrainstem(KrausandChandrasekaran,2010;StraitandKraus,2014),throughprimaryandneighborauditoryregions(Bermudez,Lerch,EvansandZatorre,2009;GaserandSchlaug,2003;Schneideretal.,2002),uptohigh-levelauditoryprocessing(Jamesetal.,2014;Loui,ZammandSchlaug,2012).Structuraldifferencesemergeatthelevelofprimaryauditorycortexandauditoryassociationareas,suchastheplanumtemporale(Bermudezetal.,2009;GaserandSchlaug2003;Keenan,Thangaraj,HalpernandSchlaug,2001;Loui,Li,HohmannandSchlaug,2011;Schlaug,Jäncke,HuangandSteinmetz,1995;Schneideretal.,2002;Zatorre,Perry,Beckett,WestburyandEvans,1998).Aparticularlypronouncedasymmetryoftheright/leftplanumtemporaleisobservedinmusicianswhoarepossessorsofabsolutepitch.Ingeneral,trainedmusiciansexhibitgreatervolumeandcorticalthicknessinauditorycortex(Heschl’sgyrus).Theseregionsaremostlikelyresponsibleforfinepitchcategorizationanddiscrimination,aswellasfortemporalprocessing.

Differencesarealsofoundatafunctionallevel,asrevealedbyfunctionalneuroimagingandneurophysiologicaltechniques(e.g.,auditoryevokedpotentials).Yet,theregionsinvolvedarenotalwaysconsistentwiththeresultsofstructuralbrainimagingstudies.Whilestructuralstudiespointtowarddifferencesinprimaryauditoryareas(GaserandSchlaug2003;Schneideretal.,2002),functionalstudiesrevealstrongerresponsesinhigher-levelauditoryregionswhencomparingmusicianswithnonmusicians.Moreover,nonmusiciansappearasneedingmoreneuronalresourcesforprocessingauditoryinformation,asshownbystrongeractivationofprimaryauditoryregionsrelativetomusicians(Besson,FaïtaandRequin,1994;Bosnyak,EatonandRoberts,2004;GaabandSchlaug2003;Shahin,Bosnyak,TrainorandRoberts,2003;Shahin,RobertsandTrainor,2004;Trainor,DesjardinsandRockel,1999).Thus,itisnotalwaysobviouswhethertrainingisassociatedwithincreasedordecreasedactivationintheunderlyingbrainregions.

Structuraldifferencesduetomusicaltrainingextendtomotorandsensorimotorcortices,topremotorandsupplementarymotorregions,andinvolvesubcorticalstructuressuchasthebasalgangliaandthecerebellum(Amunts,SchlaugandJäncke,1997;BangertandSchlaug,2006;Bermudezetal.,2009;Elbert,Pantev,Wienbruch,RockstrohandTaub,1995;GaserandSchlaug2003;Hutchinson,Lee,GaabandSchlaug,2003).Thisneuronalcircuitryisengagedinmotorcontrolandfinemotorplanning(e.g.,fingermotions)duringmusicperformanceaswellasinmotorlearning(SchmidtandLee2011).Differencesarealsoobservedintermsofbrainconnectivity(i.e.,whitematter).Forexample,musiciansexhibitgreatermidsagittalsizeofthecorpuscallosum(Lee,ChenandSchlaug,2003;Oztürk,Tasçioglu,Aktekin,KurtogluandErden,2002;Schlaug,Jäncke,Huang,StaigerandSteinmetz,1995).Thisstructure,supportingtheinteractionbetweenthetwohemispheres,maybethesubstrateofcoordinatedmovementofrightandlefthand(e.g.,fortheperformanceofcomplexbimanualmotorsequences;forinstance,seeWiesendangerandSerrien2004).Finally,theamountofmusicalpracticeisassociatedwithgreaterintegrityofthecorticospinalpathway(Bengtssonetal.,2005).

Morerecently,thereareindicationsthattrainingisnotsystematicallyassociatedwithincreasedbrainvolume.Forexample,adecreaseinstriatalvolumeisobservedinballetdancers(Hänggi,Koeneke,BezzolaandJäncke,2010)andskilledpianists(Granertetal.,2011)asafunctionofefficiencyinmotorperformance(seealsoJamesetal.,2014).Similarmixedeffectsofmusicaltrainingarefoundinfunctionalneuroimagingstudies,andarenotalwayscoherentwiththeobservedanatomicaldifferences.Musiciansexhibiteitherlowerormorelocalizedactivationintheprimarymotorcortexthannonmusiciansandmorevariablelevelsofactivationinmotorassociationregionssuchasthepremotorandsupplementarymotorareas(Haslingeretal.,2004;Hund-GeorgiadisandvonCramon,1999;Jäncke,ShahandPeters,2000;Kringsetal.,2000;Meisteretal.,2005).Theseinconsistenciesinfunctionalchangesmayreflectdifferentialrecruitmentofresourcesandmechanismsinskilledmusicians.Musicaltrainingandperformancerequiregreaterinvolvementofhigher-ordercognitiveprocesses(e.g.,tonalprocessing,workingmemory,syntax),thustheymayfavorincreaseddensityintheassociatedbrainregions.Yet,otherprocesses,

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likelytobeautomatizedandrequiringlessresourcestobeexecuted(e.g.,sensorimotorfunctionsandbasicmotorcontrol),mightrecruitlessbrainvolume(Jamesetal.,2014).Asimilarexplanationmayshedlightoninconsistenciesbetweenvisiblestructuralchangesandvariablefunctionalcorrelates(i.e.,increasedordecreasedactivationinmusicians)inprimaryauditoryandmotorregions.Thisproblemisinherentinthecomplexrelationbetweenstructuraldifferencesandfunctionalchangesassociatedwithmusicaltraining(seeZatorre,FieldsandJohansen-Berg,2012).Asystem(perceptualormotor)extendingitsrepresentationinthebrainasaresultofmusicaltrainingmayprocessinformationmoreefficientlyusingfewerneuronalresourcesthanaless-specializedsystem.Enhancedefficiencymightmanifestinlowerbloodflowdemandsinskilledmusiciansascomparedtononmusicianswhenperformingacomplexmotorsequence(Hund-GeorgiadisandvonCramon,1999;Jänckeetal.,2000;Koeneke,Lutz,WüstenbergandJäncke,2004;Kringsetal.,2000;Meisteretal.,2005).

Structuraldifferencesinthebrainofmusiciansandnonmusiciansarenotconfinedtotheauditorypathwayortomotorcircuitries.Theyextendtoparietalregions(e.g.,tothesuperiorparietallobe,includingtheintraparietalsulcusandtheinferiorlateraltemporallobe)andtotheinferiorfrontalgyrus.Althoughthefunctionalroleofthesedifferencesisstillunclear,itislikelythattheyareinvolvedinmultisensoryencodingandintegration.Sincemusicaltrainingandperformancearerichandintensemultimodalexperiencesitisexpectedthattheseactivitieswillhavebraineffectsontheinteractionbetweenmodalities(i.e.,oncross-modalintegration)(Zatorre,ChenandPenhune,2007).Notably,structuraldifferencesinfrontallobehavebeenconsistentlyreportedintheinferiorfrontalgyrus(BermudezandZatorre2005;Bermudezetal.,2009;GaserandSchlaug2003;Hanetal.,2009;Slumingetal.,2002).Ingeneral,frontalstructuresareassociatedwithavarietyofprocessesspanningfromintegrationofindividualauditoryeventsintolargerunits,mappingactionstosounds,tomusicaltonalsyntax(BangertandAltenmüller2003;Bangertetal.,2006;Koelschetal.,2002;Lahav,SaltzmanandSchlaug,2007;LevitinandMenon2003;Lotze,Scheler,Tan,BraunandBirbaumer,2003;Tillmannetal.,2006;Zatorre,ChenandPenhune,2007).Inparticulartheleftinferiorfrontalgyrusseemstoplayarelevantroleintheauditory-motornetwork.ThisregionhasbeenreferredtointhepastasBroca’sareaduetoitsfunctionsupportingaudio-motorinteractioninspeechproduction.

Functionalneuroimaginginmusiciansandnonmusiciansrevealmajordifferencesinbrainactivationslinkedtomusicaltraining,includingparietalandfrontalregions.Itisintriguingthatthesedifferencesemergealsowhenpeoplemerelylistentomusic.Forexample,priorinstrumentalpracticeisassociatedwithincreasedactivityintheauditoryandvisualsensorimotornetworkswhenperformingaperceptualtask(BangertandAltenmüller2003;Bangertetal.,2006).Thereexiststrongfunctionalconnectionslinkingauditoryperceptionandmotorprocessingduringbothmusicperceptionandperformance.Mereexposuretosoundsofwell-learnedactionstriggersaneuralnetworkextendingwellbeyondauditoryregions(BangertandAltenmüller2003;Bangertetal.,2006;D’Ausilio,Altenmüller,OlivettiBelardinelliandLotze,2006;Hasegawaetal.,2004;Haslingeretal.,2005;HaueisenandKnösche2001;Keysersetal.,2003;Kohleretal.,2002;Lahavetal.,2007;Zatorreetal.,2007).AregionwhichhasattractedparticularattentionwithinthisneuronalcircuitryisBroca’sarea,correspondingtotheposteriorinferiorfrontalgyrus.Theinferiorfrontalgyrusispartofanetworkreferredtoas“mirrorneuronsystem”(RizzolattiandCraighero2004;Rizzolatti,FadigaGalleseandFogassi,1996).MirrorneuronswereinitiallydiscoveredinregionF5inmonkeybrain,whichistreatedasthehomologofhumanBroca’sarea(RizzolattiandArbib,1998).Theinferiorfrontalgyrusisinvolvedwhenactionsareperformedbutalsowhenweseemeaningfulandgoal-directedmovements,orwhenwearepresentedwithsoundslinkedtowell-knownactions(e.g.,hearingactionwords,graspingfoodorplayingamusicalinstrument;Aziz-Zadehetal.,2006;Buccino,BinkofskiandRiggio,2004;Buccino,Vogt,etal.,2004;Lahavetal.,2007;Pulvermüller,ShtyrovandIlmoniemi,2005;Rizzolattietal.,1996).Therearesomeindicationsthatmusiciansexhibitincreasedgraymattervolumeintheinferiorfrontalgyruswhencomparedtononmusicians(Slumingetal.,2002;GaserandSchlaug2003;foraconfirmationoffunctionalinvolvementofBroca’sarea,seeSluming,Brooks,Howard,DownesandRoberts,2007),inassociationwithparietalanddorsalpremotoractivation.Thusmusicalexpertisemaybeassociatedwithgreaterinvolvementofamultimodalsensorimotor-integrationsystem,whenmusicianslistento,readthescoreof,orwatchperformancesofwell-knownmusic.

Instrument-Specific,Training-InducedPlasticity

Theevidencereviewedabovefromcross-sectionalstudiesshowsthatbrainplasticitycandifferentiatemusiciansfromnonmusicians.Notably,additionaldifferencesarefoundwhencomparingmusicianswhohavereceived

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differenttypesofinstrumentalpractice.Changesinthecorticalrepresentationswithinthemotorcortexdependontheplayedinstrument(i.e.,instrument-specificplasticity).Forexample,corticalrepresentationofthehandisdependentonthesidewhichismostinvolvedinfinemotorcontrolduringmusictrainingandperformance.Greatercorticalrepresentationsoffingersinviolinists’lefthand,ascomparedwithrighthand,havebeenfoundwithmagnetoencephalograpy(MEG)(Elbertetal.,1995).Use-specificfunctionalreorganizationofthemotorcortexisobservedwhencomparingtheshapeoftheregionscontaininghandrepresentationsinpianistsandviolinists,showinggrossanatomicaldifferencesintheprecentralgyrus(BangertandSchlaug2006).Stringplayersrequirehighlydevelopedfinemotorskillsinparticularintheirlefthand.Incontrast,inkeyboardplayers,bothhandsareassociatedwithhighlytrainedfinemotorskillswithapreferencefortherighthandasittypicallysupportsmelodyandmorearticulatedtechnicalpassageswhereasthelefthandrealizestheaccompaniment.Moreover,mostkeyboardperformersexhibitaparticularconfigurationreferredtoas“Omegasign”ontheleftmorethanontherighthemisphere,whilstmoststringplayersshowthissignonlyontheright.Theprominenceofthissigniscorrelatedwiththeageatwhichmusiciansstartedmusicalpracticeandtothecumulativeamountofpracticetime.Theobservationofthisconfigurationinrelationtothetypeofperformerarguesinfavorofastructuralplasticitymechanismdrivenbyspecificinstrumentalpractice.

Instrument-specificneuroplasticityinterestinglyextendstoperception.Musiciansshowgreaterevokedpotentialsinthepresenceofauditorystimuliascomparedtononmusicians(Pantevetal.,1998).Thiseffectismodulatedbythespecificmusicaltraining,asindicatedbytimbre-specificneuronalresponsesobservableindifferentgroupsofinstrumentalists.Forexample,stringandtrumpetplayersrevealstrongerevokedcorticalresponseswhenpresentedtothesoundoftheirrespectiveinstrument(Pantev,Roberts,Schulz,EngelienandRoss,2001),aneffectparticularlyvisibleintherightauditorycortex(Shahinetal.,2003).Inaddition,musiciansdisplayincreasedgamma-bandactivityinducedbythesoundoftheirowninstrumentascomparedtoothers(Shahin,Roberts,Chau,TrainorandMiller,2008).Thesefindingsaresupportedbyfunctionalimagingevidenceinviolinistsandflutists(Margulis,Mlsna,Uppunda,ParrishandWong,2009)indicatingthatinstrument-specificplasticityisnotrestrictedtotheprimaryauditorycortexbutratherspansacrossanetworkincludingassociationandauditory-motorintegrationareas.Recentstudiesprovideadditionalevidencethatexperience-specificplasticitymaybevisibleatthelevelofthebrainstem(Strait,Chan,AshleyandKraus,2012;forareview,Barrettetal.,2013).Insum,thereiscompellingevidenceofimportantandmeasurabledifferencesinbrainstructureandfunctionassociatedwithmusicaltrainingandlisteningexperienceinaheterogeneousgroupofmusicians.Eventhoughthesestudiesarecross-sectional,thusmakingitdifficulttoconcludeaboutacausalroleoftrainingonbraindifferences,instrument-ortimbre-specificplasticitystillsupportsthenotionofdedicatedbrainadaptations.

BeyondaCross-SectionalApproach:Short-TermEffectsofTraininginAdults

Cross-sectionalstudiesaresupportiveofstructuralandfunctionaldiscrepanciesbetweenthebrainsofmusiciansandnonmusicians.Inaddition,factorssuchasageofcommencement,intensity,ordurationofmusicaltraining/practicecanaccountfortheamountofbraindifferences(e.g.,Amuntsetal.,1997;BangertandSchlaug,2006;Elbertetal.,1995;GaserandSchlaug,2003;Schneideretal.,2002;Slumingetal.,2002).Thesefindingsandrelationspointtowardacausalrelationbetweenmusicaltraining/practiceandtheirstructuralandfunctionalbraincorrelates.Yet,themainproblemofthiscross-sectionalapproachisthatpreexistingfactorssuchaspredispositionsorgeneticcausesmayberesponsibleforsomeoftheobserveddifferencesbetweenmusiciansandnonmusicians.Longitudinalstudiesonshort-termorlong-termeffectsoftrainingareaviablealternativewhichcanallowdrawingcausalinferences.Inadults,effectsofshort-termtrainingatthebrainstemlevelarefoundinlinguisticandmusicalpitchdiscrimination(CarcagnoandPlack2011;Song,Skoe,WongandKraus,2008).Electroencephalography(EEG)andMEGstudiesrevealcorticalchanges(e.g.,increasedevokedpotentialsandincreasedsynchronizationinsecondaryauditorycortex)duetopitchdiscriminationtraining,associatedwithenhanceddetectionofpitchdiscrepancies(Bosnyaketal.,2004;Menning,RobertsandPantev,2000;Schulte,Knief,Seither-PreislerandPantev,2002),orinresponsetoimprobablesoundsfollowingrapidlearningofanovelmusicalsystem(Loui,Wu,WesselandKnight,2009).Transcranialmagneticstimulation(TMS)alsorevealschangesinhandcorticalrepresentationresultingfromshort-termtrainingofnovelfinemotorskills(e.g.,learningfive-fingerexercisesonthepiano)(Pascual-Leoneetal.,1995).However,functionalneuroimagingyieldsresultswhicharelessconsistentandmoredifficulttointerpret.Pitch-relatedlearningtasksareassociatedwitheitherdecreased(Jäncke,Gaab,Wüstenberg,ScheichandHeinze,2001;Zatorre,DelhommeauandZarate,2012)orincreasedactivation(Gaab,Gaser,andSchlaug,2006)oftheauditoryregions.Similardiscrepanciesareobserved

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followingshort-termsensorimotortraining(e.g.,piano-likeinstrumentallearning).Inthiscase,listeningtomelodiesbeforeandafterthetrainingisassociatedwitheitherincreasedactivationofregionsassociatedwithactionobservation(e.g.,thepremotorregion,Broca’sarea,andtheinferiorparietalregion)(Lahavetal.,2007),orwithdecreasedactivationofthedorsalauditoryactionpathway(Chen,RaeandWatkins,2012).Thesefindingsarereminiscentofthesituationencounteredincross-sectionalstudies,wheremusicaltrainingmanifesteditselfinincreasedordecreasedactivation.Similarly,itispossiblethatundercertaincircumstances,learningimprovesefficiencyinencodingorprocessinginformation(foranexampleinvision,seeYotsumotoetal.,2008),thusrequiringlessneuronalresources.

Traininginanexperimentalsettingcanbringaboutshort-termbrainchanges(e.g.,Chenetal.,2012;Lahavetal.,2007).Recentstudiesshowthatthenatureofthetrainingplaysarelevantroleinfosteringplasticchanges.Comparingauditoryexposurepersetoauditory-motortrainingisparticularlyinsightful.Lappeandcollaborators(Lappe,Herholz,TrainorandPantev,2008;Lappe,Trainor,HerholzandPantev,2011)submittednonmusiciansto2weeksofmusicaltraining.Inonegroup,thesensorimotortrainingconsistedoflearningtoplayamusicalsequenceonthepiano.Intheothergroup,nonmusiciansdetectederrorsinperformancesafterlisteningtothestimuliplayedbyotherparticipants.Followingthesensorimotortraining,participantsshowedenhanceddetectionofincorrectpitchortiming,ascomparedtomerelistening.Thisdifferencewasaccompaniedbylargerbrainresponse(i.e.,increasedauditorymismatchnegativity)topitchanddurationdeviations,indicatinggreaterenhancementofmusicalrepresentationintheauditorycortexfosteredbysensorimotortraining.Similarresultsshowinggreaterbenefitsofauditory-visualmultimodaltrainingascomparedtounimodaltrainingarereported(Paraskevopoulos,Kuchenbuch,HerholzandPantev,2012).Tosummarize,short-termchangesinbehaviorandbrainactivitycanbeobservedinadultsasaresultofabriefperiodofmusically-relatedtraining.Sensorimotorandmultimodaltraining,typicaloflearningamusicalinstrument,ismoreefficientindrivingneuroplasticchangesthanunimodaltraining.Thiseffectislikelytobeunderpinnedbybrainchangesoccurringbetweenauditory,motor,andsensorimotorintegrationsregions,andinvolvingbothfeedforwardmechanismscapableofpredictingtheoutcomeofmotoractivityaswellasfeedbackmechanismsformonitoringtheperformance(HerholzandZatorre2012;forexamplesinspeech,seeHickokandPoeppel2007;RauscheckerandScott2009).

MusicShapingChildBrainDevelopment

Thereisasignificantbodyofevidenceshowingstructuralandfunctionaldifferencesintheadultbrainsofmusiciansandnonmusicians.However,whetherthesedifferencesaretheresultofnatureorofnurtureisstillsubjecttodebate.Theymaybetheoutcomeoffactorswhichexistedpriortotraining(i.e.,brainpredispositionsoraptitude)orresultfrombrainadaptationsduetomusicaltrainingorexperience(e.g.,mereexposurefosteringimplicitlearning)duringsensitiveperiodsofbraindevelopment.Thereareindicationsfromcross-sectionalstudiesinadultsinfavorofexperience-dependentfactorsdrivingbrainadaptation.Earlyonsetofmusicaltraining(before7yearsofage),withinasensitiveperiod,isparticularlyefficientinstimulatingbrainchanges(forreviews,seeBarrettetal.,2013;Penhune2011;Zatorre2013;forotherdomains,suchasspeech,seeKuhl2010).Asensitiveperiodindicatesatimeframewhenearlyexperience(e.g.,musicaltraining)hasthegreatesteffectonbrainandbehaviorrelatedtotraininglaterinlife(Knudsen2004).Structuraldifferencesinthecorpuscallosumbetweenmusiciansandnonmusiciansandtheextentofhandrepresentationsinmotorcortexaregreaterformusicianswhobegantrainingbefore7years(Amuntsetal.,1997;Elbertetal.,1995;Schlaug,Jäncke,Huang,StaigerandSteinmetz,1995).Moreover,earlytrainingisassociatedwithgreaterauditorycortexandbrainstemresponsestotones(Pantevetal.,1998;Wong,Skoe,Russo,DeesandKraus,2007).Thedependenceofstructuralchangesontheageofcommencementisconfirmedwhencontrollingfortheamountoftraining(BaileyandPenhune,2010;Watanabe,Savion-LemieuxandPenhune,2007).Forexample,early-trainedmusicians(<7years)displaybettersensorimotorsynchronizationskillsascomparedtolate-trainingmusicians(>7years).Thisdifferenceisunderpinnedbybrainconnectivity(intermsofwhitematter)andstructuraldiscrepancies(intermsofgraymatter)(BaileyandPenhune,2012;Bailey,ZatorreandPenhune,2014;Steele,Bailey,ZatorreandPenhune,2013).Finally,inarecentcross-sectionalfunctionalmagneticresonanceimaging(fMRI)study,alinearregressionapproachisusedtoteaseapartage-relatedmaturationeffects,linkedmostlytofrontal(e.g.,premotor)andparietalregions,andtraining-relatedeffects,involvingmostlythesuperiortemporalgyrus(Ellisetal.,2012).

Insum,previouscross-sectionalstudiespointtowardexperience-dependentfactorsinshapingmusicians’brains.However,todeterminetherelativecontributionofnatureandnurtureinthedevelopmentofmusicalskills,

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longitudinalstudiesareneeded.Longitudinalstudiesshowbeneficialeffectsofmusicallessonsonmusicalabilitiesaswellasbehavioraleffectsonanumberofextramusicalareas.Thisphenomenonisreferredtoas“transfer”(BangerterandHeath,2004;forareview,seeKrausandChandrasekaran,2010).Neartransferisobservedwhenthetrainingdomainandthetransferdomainarehighlysimilar(e.g.,whenlearningamusicalinstrumentaffordsfinemotorskillswhichsubserveotheractivitiesbeyondmusic,suchastyping).Fartransferoccurswhenthereisrelativelylittleresemblancebetweenthetrainedabilityandthetransferdomain(e.g.,whenmusicaltrainingisassociatedwithenhancedmathematicalthinking).Fartransferofmusicaltraining,typicallymoredifficulttoachievethanneartransfer,isfoundinverbal,spatial,mathematicalthinkingandonintelligencequotient(IQ)(e.g.,Chan,Ho,andCheung,1998;Ho,CheungandChan,2003;Rauscher,ShawandKy,1993;Schellenberg2004;TierneyandKraus2013;Vaughn2000;forreviews,Schellenberg2001,2011;SchellenbergandWeiss2013).Someofthesefindings,inparticularthoserelatedtoshort-termeffectsoflimitedmusicalexposure(e.g.,theso-calledMozarteffect),arecontroversialanddifficulttoreplicate(Chabris1999;Steeleetal.,1999).Forexample,Schellenberg(2004)tested6-year-oldchildrenwhounderwentkeyboardorvoicelessonsfor36weeks.Controlgroupsofchildrenreceiveddramalessonsornolessons.Afterthetraining,childrenwhoreceivedmusiclessonsshowedasmall,albeitconsistent,increaseinfull-scaleIQandhigherperformanceinstandardizededucationalachievementtests,ascomparedtothecontrolgroups.ConsideringthatsimpleattendancetoschoolraisesIQ(CeciandWilliams1997),musicallessonsmayprovideanadditionalboostinIQbyprovidingaricheducationaloptionwhichishighlymotivatingandinvolvesawiderangeofmultimodalandsensorimotoractivities.Ataneurallevel,fartransferofmusicaltrainingmightbefosteredbysustainedinvolvementofthemultimodalsensorimotorintegrationnetworkduringrepeatedmusicallessons,drivingbrainplasticitychanges.Cross-modalplasticityinducedbymusicaltrainingislikelytoaffectregionswhicharerelevantforothertaskssuchasmathematics(parietal;Chochon,Cohen,vandeMoorteleandDehaene,1999),workingmemory(parietal;GaabandSchlaug,2003;Gaabetal.,2006),sequentialmentaloperations(frontal;Slumingetal.,2007),speech/language,auditory-motormapping,auditoryintegrationorprediction(frontal;Koelschetal.,2002;Koelsch,Fritz,Schulze,AlsopandSchlaug,2005;Lahavetal.,2007;Patel,2003;TettamantiandWeniger,2006).

Agrowingbodyofstudiesfocusesonthestructuralandfunctionalchangesoccurringinchildren’sbrainsasaresultofmusicaltraining.Insomestudies,theeffectofchildrentakingspecificinstrumentallessonswiththeSuzukimethodisexamined.Thismethod,basedontrainingtolistenbyearandlearningbyimitation,isstandardized,andthusparticularlyappropriateforsystematicstudies.Inastudy,4–6-year-oldchildrentrainedwiththeSuzukimethod(traininggroup)revealedchangesinauditoryevokedresponsestoaviolinandanoisestimulus.Inparticular,thetraininggroupshowedfasterresponsestoviolinsoundsthanthecontrolgroup(Fujioka,Ross,Kakigi,PantevandTrainor,2006;seealsoShahinetal.,2008).Notably,thesechangeswereaccompaniedbyenhancedperformanceinabehavioralmusicaltaskandimprovedworkingmemoryinanonmusicaltask.EffectsofmusicaltrainingonelectrophysiologicalbrainresponsesarenotconfinedtochildrenlearningwithSuzukimethod.Standardmusicaltrainingislinkedtogreatermismatchnegativity(MMN)responsestomelodicandrhythmicmodulationsinchildrenbetween11and13yearsofage(Putkinen,Tervaniemi,Saarikivi,deVentandHuotilainen,2014),andlargerinducedgamma-bandresponsesin5-year-oldchildren(Trainor,ShahinandRoberts,2009).Notably,enhancedbrainresponsetomusicalsoundsdoesnotrequireextensivetraining.Four-month-oldinfantsexposedtomelodiesineitherguitarormarimbatimbreforabout2.5hoursoverthecourseofaweekexhibitedMMNselectivelytothetimbreatwhichtheywereexposed(Trainor,LeeandBosnyak,2011).Furthermore,theeffectsofmusicaltrainingduringdevelopmentarenotlimitedtocorticalfunctionalitybutextendtobrainstemresponseswhenprocessingspeechinnoise(Strait,Parbery-Clark,O’ConnellandKraus,2013).Electrophysiologicalchangesareaccompaniedbyfunctionalchanges,asrevealedbyfMRI,suggestingleftwardasymmetryintask-relatedactivityduringmusicprocessing(same/differentdiscrimination),withpeaksintheauditorycortexandsupramarginalgyrus(Ellis,Bruijn,Norton,WinnerandSchlaug,2013).

Finally,musicaltrainingcanbringaboutchangesinbrainanatomy.Afterabout2.5yearsofmusicaltraining,5–7-year-old,high-practicingchildrenshowedincreasedsizeoftheanteriorpartofthecorpuscallosum,agroupoffibersconnectingmotorrelated-areasofthetwohemispheres(Schlaugetal.,2009).Thenumberofweeksofmusicalexposurewasassociatedwithchangesinthisregion,aswellaswiththeperformanceinamotor-sequencingtask.Structuralbrainchangesarealsofoundattheleveloftemporo-occipitalandinsularcortex,regionsrelatedtomusicalreading(BergmanNutley,DarkiandKlingberg,2014).Notably,braindeformationchangesinmotorandauditoryareascriticalforinstrumentalmusictrainingarevisibleassoonasafter15monthsoftrainingin6-year-oldchildren(Hydeetal.,2009).Someofthesechangesareassociatedwiththeperformance

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inbehavioraltesting.Forexample,thedeformationchangesinrightauditoryareaunderlieimprovedmelodic/rhythmicdiscrimination.Tosummarize,musicaltrainingandpracticingamusicalinstrumentareassociatedwithcognitivebenefitswhichmanifestintermsofnearorfartransfereffects.Thereisincreasingevidencepointingtothefunctionalmechanismsandstructuralchangesunderpinningthesetransfereffectsduringdevelopmentbasedonlongitudinalapproaches.Thesefindingsbegintouncovertheprocesseswhichallowmusicaltrainingtoshapefunctionalandstructuralbraindevelopment.Theeffectwouldbemostevidentwhenthetrainingisdeliveredwithinagivensensitiveperiod,whoseboundarieshavestilltobeclearlyidentified.

Conclusions

Thehumanbrainhastheremarkableabilitytochangeasafunctionoftrainingandenvironmentaldemands.Musicaltrainingandpracticearerichsensorimotorandmultimodalexperienceswhichcanshapeboththestructureandthefunctionsofhumanbrainandwhichareassociatedwithcognitivebenefits.Learningandrepeatedpracticeofaninstrumentbyprofessionalmusiciansimpliestheassociationbetweenfinemotormovementswithmeaningfulsoundpatterns,whilereceivingreal-timemultisensoryfeedback.Brainchangesaremanifestwhencomparingmusicianstononmusicians(e.g.,training-andinstrument-specificbrainplasticity),andcanbemeasuredinchildrenafterafewmonthsofmusicallyrelatedtraininginlongitudinalstudies.Behavioraleffectsandbenefitsofmusicaltrainingareundergirdedbyplasticchangesinprimaryandsecondaryauditoryandmotorcortices,aswellasinsensorimotorandmultimodalintegrationareas.

Eventhoughbrainchangesaregenerallydrivenbydedicatedmusicaltrainingasshowningroupstudies,thereareimportantindividualdifferencesinpeople’slearningofmusicalskills.Atoneendofthecontinuumthereareindividualssufferingfromlifelongdisordersofmusicallearning(i.e.,congenitalamusiaortone-deafness;forreviews,seeDallaBella,BerkowskaandSowiński,2011;PeretzandHyde2003;WilliamsonandStewart,2013).Attheotherendwefindmusicalprodigiesshowingprecociousmusicalabilities(e.g.,afinesenseofrhythmorexceptionalmotorcontrolinplayinganinstrument),whichallowthemtoachievelevelsofmusicalexpertiseunattainablefortheaveragemusician(e.g.,Kirnarskaya,2009).Indeed,thepresenceofexperience-relatedplasticitydoesnotruleoutthecontributionofpredispositionalfactors.Practiceandexperiencetypicallyoperateonnaturalabilitieswithageneticsubstrate.Thus,individualanatomicalandfunctionalpropertiesoftheneuralarchitecture(i.e.,geneticpredispositionsforlearning)mightbemoreorlessconducivetolearningmusicalskills(Zatorre,2013).Accordingly,individualdifferencesareanimportantfactorinexplainingdiscrepantresultsinneuroplasticityduetomusicaltraining(Merrettetal.,2013).Inparticular,geneticdifferenceswhichmaypavethegroundtomusicallearningaredevotedincreasingattention(Parketal.,2012;Ukkola-Vuotietal.,2013).Thereisrecentevidencethatexperience-relatedneuroplasticitycannotaccountforallobservedvariabilityinexpertbrains(FosterandZatorre,2010;Golestani,PriceandScott,2011),thusopeningthedoortogenetic/maturationalfactors.

Finally,alterationofbrainfunctionsandstructureasafunctionoftrainingorexperienceisnotconfinedtothedevelopmentalbrain,duringaparticularsensitiveperiod.Acertaindegreeofbrainplasticityisalsopossibleinadulthoodandintheelderly(e.g.,followingajugglingtraining;Boyke,Driemeyer,Gaser,BüchelandMay,2008;Draganskietal.,2004;Draganskietal.,2006).Thereareindicationsthatneuroplasticityispossibleintheadultbrainfollowingrelativelyshort-termmusicaltraining(e.g.,Herdeneretal.,2010;Lappeetal.,2008,2011).Thisfacthasmajorconsequenceswhichmotivatetheuseofmusicasatooltolimitordelaycognitivedeclineintheelderlyortoactasaneuroprotector(forareview,seeWanandSchlaug2010).Forexample,trainingprogramsrequiringintensivemultisensory,cognitive,andmotoractivities(e.g.,halfanhourofpianolessonsperweek,for6months)aresuccessfulinimprovingworkingmemory,perceptual,andmotorskills(Bugos,Perlstein,McCrae,BrophyandBedenbaugh,2007),andindelayingage-relateddeclineinspeechperception(Parbery-Clark,Anderson,HittnerandKraus,2012;Parbery-Clark,Strait,Anderson,HittnerandKraus,2011),nonverbalmemory,andexecutiveprocesses(Hanna-PladdyandMacKay,2011).Inaddition,beneficialeffectsofmusic-relatedactivitiesarefoundinstrokepatients(e.g.,Särkämöetal.,2008),andinpatientswithneurodegenerativedisorders,suchasParkinson’sdiseaseanddementia(e.g.,Limetal.,2005;Spauldingetal.,2013;VanWegen,Hirsch,HuiskampandKwakkel,2014;Vergheseetal.,2003).Understandingthemechanismsoftraining-relatedplasticityunderlyingthesebeneficialeffectsofmusicisacriticalstepfordevelopingsuccessfultheory-drivenrehabilitationstrategiesandpreventionprograms.

Acknowledgments

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ThewritingofthischapterwassupportedbyagrantfromtheEuropeanCommission(FrameworkProgramme7,ITNEBRAMUS,n.238157).

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SimoneDallaBellaSimoneDallaBella,EuroMov,MovementtoHealthLaboratory,UniversityofMontpellier-1,France;InstitutUniversitairedeFrance,Paris,France;DepartmentofCognitivePsychology,WSFiZ,Warsaw,Poland;InternationalLaboratoryforBrain,Music,andSoundResearch(BRAMS),Montreal,Canada.

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