Introduction to The Neurosciences and Music IV: Learning ... · Introduction to The Neurosciences and Music IV: Learning and Memory ... Introduction Music neuroscience research has

Ann. N.Y. Acad. Sci. ISSN 0077-8923

ANNALS OF THE NEW YORK ACADEMY OF SCIENCESIssue: The Neurosciences and Music IV: Learning and Memory

Introduction to The Neurosciences and Music IV: Learningand Memory

E. Altenmüller,1 S.M. Demorest,2 T. Fujioka,3 A.R. Halpern,4 E.E. Hannon,5 P. Loui,6

M. Majno,7 M.S. Oechslin,8 N. Osborne,9 K. Overy,9 C. Palmer,10 I. Peretz,11

P.Q. Pfordresher,12 T. Särkämö,13 C.Y. Wan,14 and R.J. Zatorre151Institute of Music Physiology and Musician’s Medicine, Hannover University of Music, Drama and Media, Hannover,Germany. 2School of Music, University of Washington, Seattle, Washington. 3Rotman Research Institute, Baycrest, Universityof Toronto, Canada. 4Department of Psychology, Bucknell University, Lewisburg, Pennsylvania. 5Department of Psychology,University of Nevada, Las Vegas, Nevada. 6Beth Israel Deaconess Medical Center and Harvard Medical School, Boston,Massachusetts. 7Fondazine Pierfranco e Luisa Mariani, Milan, Italy. 8Geneva Neuroscience Center, University of Geneva,Geneva, Switzerland. 9Institute for Music in Human and Social Development, University of Edinburgh, Edinburgh, UnitedKingdom. 10Department of Psychology, McGill University, Montreal, Canada. 11BRAMS, University of Montreal, Canada.12Department of Psychology, University at Buffalo State University of New York, Buffalo, New York. 13Institute of BehavioralSciences, University of Helsinki, Helsinki, Finland. 14Beth Israel Deaconess Medical Center and Harvard Medical School,Boston, Massachusetts. 15BRAMS, McGill University, Montreal, Canada

Address for correspondence: Katie Overy, Institute for Music in Human and Social Development (IMHSD), University ofEdinburgh, Alison House, 12 Nicolson Square, Edinburgh EH8 9DF, United Kingdom. [email protected]

The conference entitled “The Neurosciences and Music-IV: Learning and Memory” was held at the University ofEdinburgh from June 9–12, 2011, jointly hosted by the Mariani Foundation and the Institute for Music in Humanand Social Development, and involving nearly 500 international delegates. Two opening workshops, three largeand vibrant poster sessions, and nine invited symposia introduced a diverse range of recent research findings anddiscussed current research directions. Here, the proceedings are introduced by the workshop and symposia leaders ontopics including working with children, rhythm perception, language processing, cultural learning, memory, musicalimagery, neural plasticity, stroke rehabilitation, autism, and amusia. The rich diversity of the interdisciplinaryresearch presented suggests that the future of music neuroscience looks both exciting and promising, and thatimportant implications for music rehabilitation and therapy are being discovered.

Keywords: music; neuroscience; learning; memory; children

Introduction

Music neuroscience research has expanded beyondrecognition in recent years, driven not only by in-creasingly advanced, available, and affordable brainimaging technology and analysis software, but alsoby a growing interest in musical behavior within thewider disciplines of neuroscience and psychology.The field is diversifying to such an extent that whatused to be considered specialized topics can nowbe considered entire research areas, from specificaspects of music (such as rhythm or imagery) tofocused population groups (such as infants or pa-tients), to state-of-the-art techniques (such as EEG,

MEG, TMS, or MRI). Students are entering the fieldwith both undergraduate and postgraduate trainingin music psychology and music neuroscience, some-thing that could hardly have been imagined ten yearsago. There is increasing scientific and public inter-est in how music neuroscience research can poten-tially inform, and be informed by, the disciplines ofmusic therapy, music education, and music perfor-mance. The future of such research clearly capturesthe imagination and is not showing any signs ofdiminishing.

A substantial contributing factor to this success-ful expansion of the field of music neuroscience is,of course, the long-standing support of the Mariani

doi: 10.1111/j.1749-6632.2012.06474.xAnn. N.Y. Acad. Sci. 1252 (2012) 1–16 c© 2012 New York Academy of Sciences. 1

Introduction Altenmüller et al.

Foundation, a charity dedicated to child neurology.The Mariani Foundation’s pioneering conferenceshave created unique opportunities for researchersin the field, attracting students and professors alikefrom around the globe. Beginning with The Biologi-cal Foundations of Music1 in New York City in 2000,followed by The Neurosciences and Music2 in Venicein 2002, The Biological Foundations of Music: FromPerception to Performance3 in Leipzig in 2005, andThe Neurosciences and Music III: Disorders andPlasticity4 in Montreal in 2008, these internationalconferences have become key events, providing aninvaluable opportunity for meeting like-minded sci-entists, exchanging recent findings, and developingnew collaborations. The contribution to the expan-sion of music neuroscience from some key individ-uals behind these conferences (Maria Majno, LuisaLopez, and Giuliano Avanzini), along with the con-tinued support of Annals of the New York Academyof Sciences in publishing the proceedings, has beenhugely significant.

For the Neurosciences and Music-IV conference,hosted by the Institute for Music in Human andSocial Development (IMHSD) at the University ofEdinburgh in June 2011, the theme of learning andmemory was selected as the central aspect of musicalexperience. Symposia and posters were invited un-der four key topics: infants and children, adults: mu-sicians and nonmusicians, disability and aging, andtherapy and rehabilitation. Professor Richard Mor-ris gave an opening welcome to nearly 500 interna-tional delegates, and, according to tradition, this wasfollowed by an afternoon of methods workshops,this year on the topic of working with children.The following three days included a keynote lecturefrom Professor Alan Baddeley, “Human Memory;”nine platform symposia; three large and vibrantposter sessions; and a range of concerts in someof Edinburgh’s most beautiful university buildings.Here, we introduce the proceedings of The Neu-rosciences and Music IV: Learning and Memory,with each section of the volume introduced by theorganizers.

Methods I: Working with children—experimental methods

Katie OveryWorking with children is essential in order to im-prove our understanding of the development of mu-sical skills and the potential impact of early musi-

cal experiences. Learning and memory are crucialthroughout life, but particularly during early de-velopment. Noninvasive imaging techniques havebeen both difficult and rare in research with in-fants and young children, but technological ad-vances in functional magnetic resonance imaging(fMRI), electroencephalography (EEG), and mag-netoencephalography (MEG) are making such re-search easier. The design of age-appropriate behav-ioral protocols and measures are also vital, fromtask and stimuli design to child-friendly environ-ments and procedures to help discourage movementduring imaging. This collection of papers outlinescurrent experimental techniques for working withchildren, from preterm infants to kindergarten chil-dren, and includes head-turning techniques, behav-ioral protocols, fMRI, and EEG.

McMahon and Lahav5 begin with a specific dis-cussion of the problematic noise environment of aneonatal intensive care unit (NICU), which has thepotential to negatively affect preterm infant audi-tory development. After a detailed outline of thekey milestones of auditory development in utero,the authors discuss the nature of auditory plasticityat critical periods, the potential impact of being de-prived of maternal sounds after a premature birth,and evidence suggesting that noise exposure in theNICU can negatively influence the neurodevelop-ment of a child. A range of solutions is offered forreducing high noise levels and increasing exposureto natural sounds, such as offering “kangaroo care”and playing recordings of the mother’s voice.

Trainor6 continues with an in-depth discussionof current EEG and MEG methods for researchwith infants and young children. The paper beginsby explaining the source of EEG and MEG activ-ity, how they are measured, and the specific ad-vantages of these techniques when working withchildren. Trainor then describes some of the is-sues and complexities that can arise, from dif-ficulties with short attention spans and physicalmovements, to the changing morphology of wave-forms with age, to potential artifacts in the dataand how to deal with these. A series of researchexamples are provided, which show how differ-ent data analysis techniques have been used toexplore specific questions about musical develop-ment, such as the effects of exposure to differ-ent musical timbres or the development of beatperception.

2 Ann. N.Y. Acad. Sci. 1252 (2012) 1–16 c© 2012 New York Academy of Sciences.

Altenmüller et al. Introduction

Trehub7 presents a range of behavioral meth-ods for working with infants, alongside key find-ings regarding infant music perception abilities.Trehub emphasizes that the expansion of neu-roimaging technology should not replace goodbehavioral methods, which can offer important in-sights into how infants perceive and remember mu-sical information. Trehub explains the advantagesand disadvantages of different behavioral methods,regarding issues such as stimuli duration, trial num-bers, and attrition rates. Infant attention is identifiedas the key factor that must be both attracted andobserved in experimental procedures, and Trehubpoints to the benefits of ecologically valid stimulisuch as infant-directed singing. In addition, Trehubsuggests that ecologically valid infant responses areof potential importance, such as measuring rhyth-mic movements to music rather than looking be-haviors. In conclusion, Trehub advises that futureresearch in this area must use convergent measuresand should yield conceptually as well as statisticallysignificant findings.

Finally, Gaab and colleagues provide an exten-sive and invaluable survey of structural and fMRItechniques and protocols developed for infants andyoung children over the last 20 years.8 Some keychallenges are outlined, such as participant anxiety,movement restrictions in the scanner environment,and the availability of child-appropriate equipmentand pediatric brain templates. The authors go onto review a wide and detailed range of solutions tosuch challenges, from preparation sessions with amock scanner to effective data-acquisition proto-cols. A strong emphasis is placed on the need foryoung children to be comfortable, familiar with theenvironment, and engaged in the tasks presented.The ethical implications of such research are alsoconsidered, including the fact that research findingswith infants and children can potentially influencepublic policy, education, and family life.

Methods II: Working with children—social,“real world” methods

Maria Majno and Nigel OsborneA commonly held view, in many differing worldcultures and at many different times, is that the ex-perience of music has both a significant effect onan individual’s mind, body, and relationships withother human beings, and plays a useful role in thesocial, “real world.” Whereas some societies have

had systems of belief that offer an accounting forthis effect (e.g., the neo-Platonists in Europe,9 ortheorists such as Ibn Sina in the early medieval Is-lamic world10), the world of contemporary sciencehas, until recently, struggled somewhat to find con-vincing explanations and structures for reflection.

More recently, neuroscience has begun to makemajor contributions in terms of hard evidence anduseful understanding—something particularly wel-come to educationalists, music therapists, and oth-ers who seek to develop reflection on their method-ologies and to argue the case for music’s social,real-world usefulness. What has certainly helped isthat music is quite “neuroscience friendly.” It is arich, whole-brain, whole-body information that ishighly accountable. Music is communicated mostly,but not exclusively, through sound, which is a rela-tively slow mechanical energy that may be measuredscrupulously in frequencies of pitch and rhythm,precisely to the microsecond in duration, and cap-tured in Fourier transforms, spectrograms, fractals,and other representations of harmonicity, rhyth-micity, and turbulence.11 This highly accountableinformation is processed by the fastest-firing neuralsystem of the brain,12 while the mechanical energyof sound is perhaps the only energy a human beingmay “emit” in any consciously controlled, commu-nicative way.

The valuable insights neuroscience offers to mu-sic, and music’s “friendliness” to neuroscience, pointto important agendas for the future. It may be pre-mature to speak of an “applied music neuroscience,”but it is clear that many areas of social, real-worldmusical activity exist in which neuroscience may in-form practice and where practical experience mayinform neuroscience.

In the opening paper,13 Majno discusses thepsychosocial, and to some extent, political andeconomic significance of the “Sistema” initiative.The paper traces its progress from its origins inVenezuela, where ubiquitous centers provide irre-sistible musical opportunities to street children andothers (including children with disabilities) as astructured alternative to the temptation of violenceand crime, to the implementation of its methodsin Italy and other European countries (e.g., theRaploch project in Scotland14), where communitydevelopment and social organization have been pri-mary objectives. Sistema is characterized by rigor,deep learning, high standards, joy, and the premise

Ann. N.Y. Acad. Sci. 1252 (2012) 1–16 c© 2012 New York Academy of Sciences. 3


that music education should become accessible toall at no cost. The psychosocial benefits in terms ofsocial cohesion, sense of identity, and self-respectare predicated upon issues of sense of self, empathy,synchronization, emotional intelligence, memory,cognitive development, and motor skills, all close tothe domain of music neuroscience.

Uibel describes the very focused whole-child, ed-ucational, and social approach of the Musikkinder-garten Berlin,15 a project initiated by Daniel Baren-boim and intended to use music not just as an “add-on,” but rather as the central medium for day-to-daylearning. The kindergarten provides an educationin music, including a refreshing range of group cre-ative and developmental activities, and with music,including, for example, familiar partners in dramaand movement. But most radical is the work throughmusic, where music leads interdisciplinary initia-tives in areas such as topic-based learning, languagedevelopment, and numeracy.

Overy16 raises the thorny issue of negative mu-sical learning experiences and contrasts the humanmusicality of expert instrumental performance withthe more ubiquitous nonexpert human musicalityof group singing. Referring to the shared affectivemotion experience (SAME) model of emotional re-sponses to music,17,18 which emphasizes motor andsocial aspects of musical behavior, Overy proposesthat the essential and powerful nature of a musicalexperience lies not just in the sound, but also in thephysical and human origins of the sound, creating asense of shared experience. Referring to traditionalmusic education methods, Overy suggests alterna-tives to instrumental training when investigating thenature and impact of musical learning.

Osborne considers the influence of developmentsin neuroscience on reflection on practice in the useof music to support children who are victims ofconflict.19 This paper locates the work within an en-hanced bio-psycho-social framework. Although thestandard diagnostic instruments for PTSD (post-traumatic stress disorder) are concerned with is-sues such as traumatic events, traumatic recall,avoidance, and hypervigilance, there are signifi-cant physiological symptoms, such as raised heartrate, respiratory problems, hyperactive or sluggishmovement repertoires, and the dysregulation ofendocrine systems (including the hypothalamic–pituitary–adrenal axis) and neurotransmission. Os-borne further traces a circle around the bio-psycho-

social framework to include aspects of communica-tive musicality, emotional communication, empa-thy, self-belief, creativity, and identity, relating eachto emerging research in neuroscience. It is impor-tant for any music intervention to avoid inappro-priate invasion of the physical and mental lives oftraumatized children. But evidence from work inneuroscience and related disciplines with nontrau-matized populations has been applied to supportcritical reflection on experimental methodologiesseeking to use music to help regulate, for example,motor activity, the autonomic nervous system, andendocrine and respiratory systems.20

Symposium 1: Mechanisms of rhythm andmeter learning over the life span

Erin HannonDancing and moving in time with music are uni-versal capacities that constitute a crucial compo-nent of human musical experience. Growing ev-idence suggests that rhythm and beat perceptionrely on integration of information across multiplesensory modalities and a broad network of brainregions.21–23 A fundamental goal of research onrhythm and meter perception is to understand theconditions under which a beat or hierarchical met-rical structure can be inferred from a given stimu-lus, the mechanisms underlying perception of a beator meter, and the extent to which metrical percep-tion depends on casual listening experience, formalmusic training, or simply emerges from the interac-tion between the brain and the stimulus itself. Thissymposium attempted to tackle these fundamentalquestions using various methodological approachesand technologies, and testing individuals of differ-ent ages and/or abilities.

McAuley24 uses fMRI to examine neural corre-lates of beat perception. The study uses a sequence-timing paradigm that typically produces large indi-vidual differences in sensitivity to the implied beat;however, beat sensitivity declines as the stimulus ispresented at slower and slower tempos. By com-paring brain responses under conditions of highand low beat sensitivity, the study provides evi-dence for specific brain networks that are associatedwith beat- versus interval-based temporal process-ing. Honing25 tackles the question of whether hi-erarchical metrical representations arise automat-ically in the brain or require extensive listeningexperience or formal music training. The paper



provides a tutorial on developing optimal rhyth-mic stimuli and tasks that may demonstrate hierar-chical beat perception through EEG by measuringa mismatch-negativity (MMN) response to stimu-lus omissions. It reviews evidence from EEG studieswith adults and newborn infants that suggest hierar-chical beat perception may be possible with minimalprior experience or training. Hannon, der Nederlan-den, and Tichko 26 also explore the role of experiencein meter and beat perception. They use similarityjudgments to examine how well American childrenand adults detect beat-disrupting changes to famil-iar (Western) simple-meter and foreign (Balkan)complex-meter folk songs before and after at-homeexposure to foreign complex-meter music. The find-ings indicate that listening exposure dramaticallychanges how children under age 10 perform in thejudgment task, with accuracy improving for com-plex meters but declining for simple meters. Bycontrast, adults and older children consistently per-form more accurately in the simple- than in thecomplex-meter conditions, and this trend is vir-tually unchanged by at-home listening experience.This suggests that at least some of the metrical rep-resentations that contribute to beat perception un-dergo slow developmental change over the courseof childhood and that listening experience can fine-tune these representations during early but not latechildhood or adulthood.

Together, the three papers raise important ques-tions about the nature of beat-based perception.On the one hand, rhythmic patterns appear to ac-tivate beat-based expectancies even in highly inex-perienced newborn listeners.25 On the other hand,these expectancies are obviously influenced by lis-tening experience and presumably by the acquisi-tion of culture-specific musical knowledge.26 As re-search continues to illuminate our understandingof the neural underpinnings of beat-based process-ing,24,25 it may be possible to disentangle the var-ious processes and mechanisms that contribute toour subjective experiences of beat and meter.

Symposium 2: Impact of musical expertiseon cerebral language processing

Mathias S. OechslinIn cognitive neuroscience, the question of whethermusical expertise affects or facilitates language pro-cessing has been increasingly addressed. In this sym-posium, we introduced new studies that observe

the influence of musical expertise on various as-pects of neural language processing—making use ofseveral brain imaging and cognitive measurements,such as event-related brain potentials (ERP), fMRI,and behavioral performance data. This spectrum ofmethodological approaches characterized by bothtime and spatial sensitivity enables a deeper under-standing of musicians’ neural processing of fine-grained acoustic language properties. Together, thepapers in this section provide insight into languageprocessing, ranging from basic auditory segmen-tal speech information to suprasegmental decodingmechanisms at word and sentence level.

Disclosing cortical plasticity by focusing on basalauditory functioning, previous studies have un-veiled that musical expertise alters structure andfunctional mechanisms of various brain areas thatare involved in processing of both speech and mu-sic.27–30 Therefore, it is reasonable to assume thatneural encoding in the former domain could ben-efit from the expertise in the latter—even thoughpercepts of speech and music are phenomenologi-cally completely distinct.

Regarding basal auditory functioning, two re-cent studies31,32 have demonstrated preliminary butcompelling evidence that musical training inducedplasticity in segmental speech processing. Perform-ing an auditory (phoneme discrimination) cate-gorization task, musicians and nonmusicians wereasked to identify voiced and unvoiced consonant–vowel (CV) syllables with either natural or broad-band noise acoustic spectra. The fMRI study re-vealed higher accuracy in identifying CV-syllableswith manipulated spectra, flanked by a leftwardhemispheric asymmetry and overall enhanced acti-vation of the planum temporale in musicians com-pared to nonmusicians.32 Moreover, using the sameexperimental paradigm, electrophysiological datarevealed that N1 amplitudes and corresponding to-pography were identical in voiced and unvoicedstimuli in musicians but not in nonmusicians. Ac-cordingly, in their paper, Meyer et al. comprehen-sively discuss the role of the planum temporale forspeech processing as a function of musical exper-tise.33 Taken together, these results suggest that mu-sical expertise leads to a honed auditory capacity inspectrotemporal analysis that goes far beyond thedomain of music. In this context, Besson et al.34

have argued that “enhanced sensitivity to acousticfeatures that are common to music and speech, and



that imply domain-general processes, allow mu-sicians to construct more elaborated percepts ofspeech processing than nonmusicians. . .[and] fa-cilitates stages of speech processing that are speech-specific.”

Moving on along this line to the processing oflinguistic structures (word segmentation level), theinfluence or interaction of such sensory plasticitywith expertise-dependent top–down modulations(see later) is not yet clear and should be observedin more detail. However, it has been demonstratedthat the nature of implicit knowledge in music andlanguage, in general,35 and learning of linguisticstructures and musical structures, in particular, areclosely related:36 by analyzing behavioral data of astatistical learning paradigm, performance in a lin-guistic test has been found to be positively correlatedwith performances in a musical test (based on con-catenated results of Refs. 37 and 38). In the latterERP study,38 musicians and nonmusicians were ini-tially presented with sequences of artificial (sung)language. Then participants were asked to make fa-miliarity ratings of musical and speech sound pairs,indicating whether the first or the second item re-sembled the previously presented sung sequence.Most interestingly, Francois and Schön found en-hanced N400 amplitudes in musicians compared tononmusicians, elicited by both the linguistic andthe music task, concluding for musicians an advan-tage in stream segmentation or in lexical storage orboth. The authors of this paper provide valuableinsight into technical details of statistical learningparadigms.39

In a recent publication, Patel40 systematically ad-dressed the prerequisites of such possible transfereffects from musical training to neural encoding ofspeech. In what is formulated as the OPERA hypoth-esis, Patel concluded that overlapping brain net-works processing shared acoustic features in speechand music are the ultimate essentials for possibletransfer effects. Furthermore, to end up with a ben-efit in speech encoding, according to Patel, threeadditional conditions have to be fulfilled, includingaspects of musical processing demands, training in-tensity, and emotional arousal. Here, Patel presentshis most recent view featuring the fundamental as-pects of different precision demands in music andspeech processing.41 Originally, the OPERA hypoth-esis mainly referenced studies that observe musicaltraining-induced subcortical plasticity driven by de-

scending auditory projections.42 By analyzing suchbrainstem responses, it has been revealed that mu-sical expertise yields enhanced auditory frequencyrepresentation,43 increased accuracy in encodinglinguistic pitch patterns,44 and facilitation of hearingspeech in noise.45 Moreover, the latter acuity is sup-ported by cognitive mechanisms, such as enhancedtonal working memory45 and auditory attention46

in musicians compared to nonmusicians. Accord-ingly, topographical ERP analyses confirmed thatprefrontal responses are considerably altered (lowerprefrontal auditory response variability) as a func-tion of musical expertise during a selective auditoryattention task.46 In other words, this result signi-fies advantages for musical experts in situations thatrequire sustained auditory attention. In their paper,Kraus et al. specifically focus on musicians’ auditoryadvantages and the mediating role of auditory work-ing memory.47 In fact, previous fMRI research foundthat attention driven top–down modulations facili-tate basal auditory speech processing.48 This frame-work lucidly implies that future research is neededto disentangle the potential influence of musical ex-pertise on interactions among structural brain char-acteristics, cognitive functions, and the architectureof cerebral language processing.

This ensemble of studies that addresses the sametopic from different directions is clearly indicative ofthe increasing relevance of research on transfer ef-fects from music to language mechanisms at severallevels of complexity.

Symposium 3: Cultural neuroscienceof music

Steven M. DemorestThe goal of the study of the cognitive neuroscienceof music is to explain how brain structure and func-tion interact with and shape musical thought andbehavior. Research in this area has been dominatedby Western thought regarding the nature of musicperception and performance, a bias that has the po-tential to limit the scope of our theories regardingmusic and brain function. The implicit learning ofone’s culture or enculturation seems to affect virtu-ally every area of human thought, and the relativelynew field of cultural neuroscience49 is beginningto document the important influence of culture inshaping brain function. Given that music is one ofthe primary agents of cultural transmission, it standsto reason that certain aspects of musical thinking are



mediated by culture. This symposium explored cur-rent research in the cultural neuroscience of musicand how findings in this field might clarify the roleof culture in music development, cognition, andlearning.

Previous research in infant music perceptionhas identified that encultured responses to musicemerge at around 12 months of age.50 Here, Trainoret al. document how early musical experiences canhasten culturally biased responses in infants.51 In aseries of experiments, they explored the effect of ac-tive participation in Kindermusik classes and otherformal music exposures that influence infants’ be-havioral and brain responses to Western rhythm,timbre, and tonality. Vuust et al. also present datashowing learning-based changes in brain responses,but in adult populations,52 they used an innovativeMMN paradigm to differentiate the responses ofperformers from different genres of Western musicto acoustic changes in a complex musical stimu-lus. Tervaniemi and her coworkers extend researchon classical musicians’ ERP responses to harmonicviolations53 to Finnish folk musicians,54 whose re-sponses reflect highly specialized sensitization toharmonic violations. Large and Almonte proposea new theory of tonality that seeks to uncover thebasic neurodynamic principles that underlie tonalcognition and perception. They provide evidencethat auditory neurons respond to tonal relation-ships in predictable ways and discuss the possibilityof general neurodynamic principles that underlie alltonal cognition.55

Demorest and Wong both explore how implicitlearning of culture influences adults’ neurologicalresponses to music of different cultures using ERPand fMRI methodologies, respectively. The first pa-per explores crosscultural music cognition by track-ing ERP responses to melodic expectancy viola-tions in culturally familiar and unfamiliar music.56

As in previous research,57 Western listeners exhib-ited a P600 response to scale deviations in West-ern melodies. They exhibited a smaller P600 whenhearing similar deviations in north Indian classicalmelodies, which may reflect a cultural general sen-sitivity to statistical properties of tonal sequencesor possible areas of overlap between the two tonalsystems under study. In the second paper, Wonget al. explore fMRI responses to affective judgmentsacross two musical cultures and how they differ be-tween monomusicals and bimusicals.58 In addition

to confirming earlier findings of different behav-ioral responses,59 they found significant group dif-ferences in the connectivity of a temporal–limbicnetwork. This provides evidence of qualitatively dif-ferent brain function as a result of exposure to twoculturally distinct musical systems. The papers pre-sented here touch on some of the central issues incultural neuroscience, including the interaction ofenculturation, formal training, and development.

Symposium 4: Memory and learningin music performance

Caroline Palmer and Peter Q. PfordresherThe study of music performance has important im-plications for understanding basic mechanisms ofmemory and learning. In contrast to listeners, whoacquire vast implicit knowledge resources accumu-lated during years of perceptual experience, individ-uals differ greatly in the explicit and implicit knowl-edge they have acquired for performance. Theseindividual differences in performance knowledgeleave open the possibility for studying the develop-ment of performance-specific learning and memoryin cross-sectional and longitudinal contexts, withchild and adult populations. In addition, music per-formance typically requires the execution of long,rapid sequences typically performed from memory,thus providing a rich context in which to study thedevelopment of sensorimotor learning, as perform-ers learn to map auditory and visual feedback tospecific actions. Finally, disorders that affect per-formers offer an excellent venue in which to studymusic’s role in rehabilitation from sensory and mo-tor disorders.

The first paper in this section, by Bailey and Pen-hune,60 explores the acquisition of musical skillsat different points in the life span to determinewhether this is a “sensitive period” for musicalskill acquisition. In keeping with the assumptionof a sensitive period for musical development, adultmusicians who began training early in life exhib-ited better synchronization ability than adult non-musicians or musicians who began training laterin life. Music performance is fundamentally inte-grative, involving the online coordination of ac-tions with the perception of feedback from thoseactions.

The next two papers consider the role of per-ceptual feedback in performance. Pfordresher ad-dresses how musical training influences the role of



auditory feedback in nonmusicians.61 Contrary toa purely associationist view, nonmusicians do notapproach the novel task of music performance as“blank slates” but instead build on preexisting sen-sorimotor associations, based on their broader ex-perience of perception/action associations, that arerefined through musical training. Zimmerman andLahav’s review62 focuses on the neural underpin-nings of multisensory associations that are enhancedby musical training. They argue that the multisen-sory processing that is enhanced by music listeningand performance underlies the success that music-related therapy has had for rehabilitation.

The fourth paper, by Palmer et al.,63 discusses thetime course of retrieval failures and their sensori-motor consequences during performance. They re-port the results of an experiment in which pianistspracticed and performed novel musical pieces atfast and slow tempi. Both pitch errors and the cor-rectly produced events immediately prior to thoseerrors were produced with lower intensities and re-duced tempo from other correctly produced tones,consistent with the view that performers covertlymonitor for production errors prior to responseselection.

The final paper, by Felix Strübing and MariaHerrojo Ruiz,64 examines how sensorimotor pre-diction is implemented in healthy and dystonic pi-anists by recording neural responses to errors usingevent-related potentials (ERPs). Results suggesteddegraded error processing among dystonic patients,even preceding error production. Together, thesepapers intersect with the developmental, skill acqui-sition, and rehabilitory themes of the other papersin this volume to elucidate the neural foundationsof memory for musical experience.

Symposium 5: Mind and brain in musicalimagery

Andrea R. Halpern and Robert J. ZatorreMental imagery refers to a type of memory repre-sentation that is rich in perceptual detail. For in-stance, when asked to describe one’s living room,most people can report the spatial layout, colors ofthe walls, and textures of the furniture. Auditoryimages also contain perceptual information. Thereader is invited to imagine a piece of familiar music,perhaps “Happy Birthday.” Musicians and nonmu-sicians alike can report auditory characteristics oftunes, such as the pitch, rhythm, and tempo.65 Thus

the experience can be quite vivid, sometimes so vividthat the auditory experience takes the form of a per-sistent repetitive memory, colloquially referred to as“earworms.”66 In addition to vividness, in severalmeaningful ways auditory imagery for music canalso be said to capture qualities of the actual pieceand is thus a veridical representation. For instance,people are willing to set a metronome to the tempoof an imagined familiar tune, or indicate on a key-board the starting pitch they habitually assign to thetune, which they will replicate reliably over multi-ple trials.67,68 People without absolute pitch abilitiescan fairly accurately report the opening pitch or keyof familiar recorded music.69,70

In addition to behavioral research, a number ofstudies have documented the similarity of neuralmechanisms in imagery and perception. In a priorreview,71 Zatorre and Halpern examined the sub-stantial evidence that the secondary auditory cortexis active when people imagine music, again suggest-ing that we co-opt perceptual mechanisms whenretrieving these vivid and veridical experiences. Thereview also noted that these experiences are not con-fusable, and that many studies find additional acti-vation in the frontal cortex during imagery that isabsent in perception, suggesting an important mod-erating influence of executive and memory func-tions on the auditory cortex.

This symposium at the conference examined newresearch in auditory imagery that extends earlierstudies on basic aspects of auditory imagery. Amongthe four speakers, a number of methodological ap-proaches were represented: behavioral studies, com-puter simulations, measurement of electrical activ-ity from the brain surface (ERP), and measurementof relative blood oxygenation as a proxy for neuralactivity (fMRI). The variety of approaches remindsus of the importance of both “mind” and “brain” inour symposium title.

Three types of extensions to earlier work wereevident among our speakers. First, several speakerspresented tasks that required experimental partic-ipants to make rather fine distinctions during im-agery tasks. This is perhaps most evident in Janata’spaper on the accuracy with which people can makein-tune and out-of-tune judgments in imaginedtones.72 Halpern reports here on the ability to makemoment-to-moment judgments on emotional as-pects of imagined music using a continuous scale,73

and Zatorre reports on neural correlates of subtle



key-distance effects during mental transformationsof music.74

Second, many of the studies discussed requiredfairly complex processing, particularly regarding theextent to which working memory was required forsuccessful completion. As one example, Halpernreports a study in which people learned pairs oftunes and were required to anticipate the secondpair member upon presentation of the first.73 Kellerreports studies of several aspects of musical perfor-mance that need to be maintained in parallel forsuccessful execution.75 Arguably, the most difficultstudy in the set, reported by Zatorre, was askingpeople to imagine tunes in reverse note order!74

This imposes a very high working memory loadand was reflected in extensive frontal and parietalactivations.

Third, several speakers framed their tasks inthe context of the adaptive usefulness of auditoryimagery. The most obvious example of this wasKeller’s exploration of the role of auditory imageryin synchronization of movements among musicalperformers.75 Janata also suggested that auditoryimagery actually enhances the fine pitch judgmentsrequired in the tasks he presented, and Halpernnoted that the ability to extract emotion from imag-ined music adds to the enjoyment of rememberedexperience.73

Overall, the talks in the symposium showed thatconsiderable progress has been made in extendingthe breadth and depth of knowledge about the qual-ity and richness of the musical sounds we hear andmanipulate in our minds, despite the essentially pri-vate nature of mental imagery.

Symposium 6: Music-induced adaptiveand maladaptive brain plasticity in healthand disease

Eckart AltenmüllerEmerging research over the last decade has shownthat long-term music training and the associatedsensorimotor skill learning can be a strong stim-ulant for neuroplastic changes both in the devel-oping and adult brains, affecting both white andgray matter as well as cortical and subcortical brainstructures.76 Making music, including singing anddancing, leads to a strong coupling of perception andaction mediated by sensory, motor, and multimodalbrain regions and affects, either in a top–down orbottom–up fashion, important relay stations in the

brainstem and thalamus.77 Furthermore, listeningto music and making music provokes motions andemotions, increases between-subject communica-tions and interactions, and is experienced as a joy-ous and rewarding activity through activity changesin the amygdala, ventral striatum, and other com-ponents of the limbic system.78 Music is a powerfultool in rehabilitation, providing an alternative entrypoint into compromised neural circuits due to braindamage.79 Music thus can remediate impaired neu-ral processes or neural connections by engaging andlinking brain regions with each other that mightotherwise not be linked together. The pleasurablepower of listening to favorite music can even reversemaladaptive changes in the auditory cortex, caus-ing the torturing tinnitus percept.80 On the otherhand, music-induced brain plasticity has its darksides. Prolonged practice, high workload, overuse,and extreme demands on sensorimotor skills in pro-fessional musicians may result in a degradation ofexactly these fine motor abilities, a condition termedmusician’s dystonia.81

The aim of this symposium was to summarizethe latest research concerning the powerful impactof music on brain plasticity in both directions—adaptive and maladaptive. In the first article, Schulzeand Koelsch present a review of behavioral andneuroimaging findings on similarities and differ-ences between verbal and tonal working memory inmusicians and nonmusicians.82 They demonstratethe impact of musical training on verbal memoryand its consequences for verbal learning in chil-dren and adolescents. Novel results are discussedthat imply the existence of a tonal and a phono-logical loop in musicians, based on partly differ-ing neural networks underlying verbal and tonalworking memory. Finally, the authors propose thatboth verbal and tonal auditory working memory arebased on the knowledge of how to produce the to-be-remembered sounds, and therefore sensorimo-tor representations are involved in the temporarymaintenance of auditory information in workingmemory.

In the second article, Pantev presents a noveland promising method to reduce chronic tonaltinnitus.83 There is evidence that maladaptive au-ditory cortex reorganization may contribute to thegeneration and maintenance of this torturing con-dition, which can be conceived as a permanentcortical memory trace. Because behavioral training



can modify cortical organization, an approach waschosen to expose chronic tinnitus patients to self-chosen, enjoyable music, which was manipulated tocontain no energy in the frequency range of the indi-vidual tinnitus, thus promoting lateral inhibition tothe brain area generating tinnitus and overwritingthe dysfunctional cortical memory.

The third article, authored by Zipse et al. from thelaboratory of Schlaug, presents an impressive casereport on rehabilitation of an adolescent stroke pa-tient suffering from Broca’s aphasia.84 Using a mod-ified version of melodic intonation therapy (MIT),the behavioral improvements were accompanied byfunctional MRI changes, demonstrating that theright frontal lobe takes over language functions. Thiscase study not only provides further evidence for theeffectiveness of this rehabilitation strategy, but alsoindicates that intensive treatment can induce func-tional and structural changes in a right hemispherefrontotemporal network.

Jäncke presents in his article on the dynamicaudio-motor system in pianists a pilot study basedon the theoretical assumption that continuousclosed-loop audio–motor control could be disad-vantageous for pianists.85 He argues that the func-tional relationship between the intracerebral electri-cal activations in the auditory and premotor cortexshould be rhythmically decreased and increased. Totest this hypothesis, activations and connectivity ofthe auditory and premotor cortices were estimatedusing a novel method to analyze EEG time seriesand their causal relationship, similar to the “dy-namic causal modeling” approach used in fMRI.The analysis revealed a “causal relationship” fromthe auditory cortex to the premotor cortex, whichwas considerably stronger during piano playing andweaker during rest. Interestingly, this relationshipvaried rhythmically during the course of piano play-ing, thus delivering evidence that in professional pi-anists, the functional coupling between the auditoryand premotor cortex is instable, highly dynamic, andextremely adaptive.

Finally, Altenmüller et al. focus on musician’s dys-tonia as a syndrome of dysfunctional brain plas-ticity.86 This condition is characterized by a lossof fine motor control of extensively practiced andhighly skilled movement patterns in professionalmusicians. On a neurophysiological level, this phe-nomenon is due to fusion of sensorimotor recep-tive fields in the cerebral cortex and to a lack of

lateral inhibition to adjacent body parts. In theirpaper, the authors demonstrate that behavioral fac-tors can trigger the manifestation of this disablingdisorder. The interplay between genetic predisposi-tion, psychological and behavioral factors, such asperfectionism, anxiety, and over-specialization, pre-dominantly in classical reproductive musicians, iselucidated, and its important role in finally causingthis neurological condition is convincingly demon-strated.

Symposium 7: The role of music in strokerehabilitation—neural mechanisms andtherapeutic techniques

Takako Fujioka and Teppo SärkämöNeuroplasticity is a key mechanism underlyinglearning new skills and relearning lost skills dur-ing rehabilitation. Recent cognitive neuroscienceresearch has shown that training in music mak-ing can enhance neural processing in sensory, mo-tor, executive, and affective brain systems and fa-cilitate interactions between those brain systemsin healthy children and adults.87–90 This is likelydue to the fact that the massed practice associatedwith music making requires enormous resourceswithin each specific brain system as well as ex-tensive coordination between the systems. Impor-tantly, music making also involves social interac-tion and induces strong emotional experiences. Evensimply listening to music can have a short-termpositive effect on arousal, attention, memory, andmood.91–93

The next question is how these findings can beincorporated and translated into the rehabilitationof the damaged brain. One of our strong allies inanswering this question is the music therapy com-munity, which has accumulated and practiced theo-ries on how music can be used to help neurological,psychological, and functional recovery in variousclinical populations, including stroke patients. An-other guiding principle is the knowledge obtainedfrom rehabilitation sciences that meaningful andrelevant tasks and motivated participation in activ-ities enhance rehabilitation outcomes in everydaylife, compared to mere repetition of rote exercises.Thus, it seems that we have reached the crucial pointat which each discipline can help inform the othersin ascertaining how music can be used for improv-ing the lives of patients.



How can music-related activities fit into rehabil-itation? Within the framework of the InternationalClassification of Functioning, Disability and Health(ICF) of the World Health Organization (WHO),94

we propose that music making and/or music listen-ing activities can be understood as an activity thatinfluences a stroke patient’s environment, which inturn interacts with behavioral activity and partici-pation, as well at the level of brain function. For ex-ample, a music-making exercise demands not onlysensorimotor but also auditory memory functionsincorporated with motor planning and execution.Increasing complexity and challenge can easily bebuilt into musical exercises, such that they are inline with the principles of effective motor learning.Also, the goal-oriented approach of music makingresonates with the importance of meaningful tasksin rehabilitation. Compared with many traditionalmotor and cognitive rehabilitation methods, music-based activities have the advantage of being intrinsi-cally motivating and fun as well as providing directand natural feedback, which can be crucial for en-gaging patients in their rehabilitation process. Atthe same time, engagement in music making canbecome an important emotional event, often pro-viding solace and comfort and relieving stress andanxiety, and can therefore have an important addi-tional positive impact on quality of life.

In this section, we synthesize recent studies ofmusical interventions for stroke survivors. The in-terventions are based on existing evidence regardingunderlying brain mechanisms, and outcome mea-sures are demonstrated as functional changes at thelevel of behavior and the brain. These give us impor-tant insights into how the human brain in both thehealthy and damaged state processes music, withits multimodal interactions. In the first article,95

Särkämö and Soto review current evidence of theeffects of music listening on emotion, cognition,and the brain; present two bodies of experimentalstudies showing that listening to pleasant music aftera stroke can temporarily enhance visual awarenessin patients suffering from unilateral spatial neglectas well as improve the recovery of memory, atten-tion, and mood; and discuss the potential neuralmechanisms underlying the rehabilitative effect ofmusic listening. In the second article,96 Rodriguez-Fornells, Rojo, Amengual, Ripolles, Altenmüller,and Münte review the literature concerning the

application of music-supported therapy (MST) (anew motor rehabilitation method that uses musi-cal instruments) and present experimental resultsindicating that MST is effective for improving therecovery of motor skills and mood after a stroke,with a direct impact on the activity and functionalconnectivity of the frontotemporal auditory–motornetworks. MST is also discussed in the third ar-ticle97 by Fujioka, Ween, Jamali, Stuss, and Ross,which demonstrates, using MEG, that listening andtapping to a beat are both associated with the pe-riodic modulation of beta oscillations in the brainthat are typically linked to motor functions, andthat MST can induce changes in the contribution ofauditory and motor brain areas to the beta activityafter chronic stroke. Their data also give some in-sight into how rhythmic auditory stimulation (RAS)using music or metronome sounds may successfullyenhance rehabilitation exercises through activatingthe sensorimotor beta-band network.98,99

In the fourth article,100 van Wijck et al. intro-duce the rationale and methodological basis of anovel, innovative motor intervention for stroke pa-tients, which integrates the patient’s preferred mu-sic with game technology in a rhythmic auditorycueing task for training upper limb function. Fi-nally, Tomaino101 describes various singing-relatedtechniques used in clinical music therapy to en-hance speech ability in nonfluent aphasic patients.Although MIT has been known as an effectivesinging exercise,84,102 the paper documents how di-verse the symptoms of aphasic patients can be andhow emphasizing different musical features and so-cial/motor cues in the practice can help with treatingthem differently.

In summary, this section highlights recent ad-vances in the field of music therapy for stroke andprovides steps toward integrating musical activitiesinto the neurological rehabilitation of stroke pa-tients and toward understanding how music canwork in the damaged brain. Although the findingsare encouraging, the field is still new, and more re-search is clearly needed for building a solid evidencebase through randomized control trials that com-pare musical interventions against adequate controlinterventions. Through this synthesis, our hope isto inspire and stimulate future research directionswith joint efforts among neuroscientists, therapists,psychologists, and engineers.



Symposium 8: Autism and music

Catherine Y. WanAutism spectrum disorder (ASD) is a developmen-tal condition that affects one in 110 children, andas many as one in 70 boys.103 In addition to so-cial abnormalities and the presence of repetitiveand stereotyped behaviors, one of the core diag-nostic features of ASD is impairment of languageand communication.104 Severe deficits in commu-nication not only diminish quality of life for affectedindividuals but also present a lifelong challenge fortheir families.

Despite their social and communication deficits,children with ASD can enjoy auditory–motor activ-ities, such as making music, through singing or play-ing an instrument. In addition, such children oftendisplay enhanced music and auditory-perceptionabilities.105 In the first published report of autism,Kanner106 described the exceptional musical skills ofseveral children, including one notable example ofan 18-month-old boy who was able to discriminateamong many symphonic pieces of music. Recentinvestigations have provided further evidence forenhanced music-perception abilities in individualswith ASD. Often described as emotionally unreach-able or in their own world, these individuals arenevertheless often able to express typical affectiveresponses to music and to derive enjoyment frommusic.

The paper entitled “Music: a unique window intothe world of autism” presents recent advances inresearch examining neurobiological underpinningsof musical processing in ASD and the therapeuticpotential of music making in ameliorating some ofthe associated deficits. Molnar-Szakacs and Heatonsummarize research on the dissociation betweenemotional communication abilities in the musicaland social domains in individuals with ASD.107 Al-though individuals with ASD are often impaired intheir ability to understand nonverbal expression ofemotions, they can nevertheless understand simpleand complex musical emotions.17 Potential expla-nations for such uneven development across func-tional domains, as well as future directions for thestudy of music in autism, are discussed.

Hyde and colleagues review the behavioral andneuroimaging literature on auditory pitch and timeprocessing in ASD.108 Individuals with ASD gen-erally show enhanced perceptual skills in a musi-cal context but not in a linguistic context. These

observations may be related to the locally ori-ented or highly focused behaviors that are com-mon in ASD.109 Identifying the brain–behavior re-lationships of auditory processing in ASD may helpto identify neurobiological markers and enhancedtreatment potential in ASD.

Wan et al. present diffusion tensor imaging datacollected from a group of completely nonverbal chil-dren with ASD.110 Abnormalities in a language-related white matter tract, the arcuate fasciculus,were investigated. This tract connects auditory andmotor brain regions and is important in the map-ping of sounds to articulatory actions during speech.The preliminary imaging findings reported heremay explain why children with ASD fail to developspeech naturally. Furthermore, the findings com-plement the laboratory’s ongoing treatment studyof a novel intonation-based intervention (auditory–motor mapping training, AMMT), which aims tofacilitate speech output in nonverbal children withASD.111 It is suggested that interventions that aredesigned to engage abnormal auditory–motor con-nections (such as AMMT) have the potential toeffectively facilitate the development of expressivelanguage skills.

Taken together, the three papers on music andautism review current knowledge in this emergingfield, which will hopefully stimulate further researchand the development of interventions for a disorderthat affects the lives of a great many individuals.

Symposium 9: Learning and memoryin musical disorders

Psyche Loui and Isabelle PeretzAlthough music is ubiquitous across human cul-tures, a subset of the normal population appears tohave an abnormal lack of musical ability. Increasingevidence from behavioral, neuroimaging, and ge-netic studies has documented difficulties in pitchperception as well as production, also known ascongenital amusia or tone deafness.112,113 Althoughthese substantial deviations from the musical normare fascinating to the general public, they can alsobe informative as a window into the psychologicaland neural capacities necessary for musical func-tioning. Furthermore, they can provide us with amodel system through which to investigate capac-ities that, like music, might be uniquely human,such as speech and language, expectation for futureevents, and consciousness.



Given that musical disorders may have impor-tant implications for a broader scientific commu-nity, there is a need to have a solid understandingof their underlying cognitive processes and mecha-nisms. How do musical disorders inform our modelof the human capacity for music? To date, testedtheories of the causes of musical disorders includedifficulties in fine-grained pitch discrimination,114

spatial processing115 (see also Ref. 116), pitch aware-ness,117 disconnection of neural pathways,118 andshort-term memory limitations in pitch.119 Despitethese theories, we know relatively little about howcognitive constraints might limit musical perfor-mance. In particular, limitations on memory areonly recently becoming addressed as possible expla-nations for musical disorders. Given that personswith congenital amusia have pitch-specific limita-tions on their short-term memory resources thatare independent of perceptual difficulties,120 it re-mains to be seen whether and to what extent thesememory limitations could give rise to learning diffi-culties. If learning is indeed affected in people withmusical disorders, then to characterize these learn-ing deficits, and possibly to design rehabilitationstrategies, we need to know about the types of in-formation learning that are affected: whether theselearning difficulties are circumscribed to music, andwhether related modalities such as language learn-ing might be affected as well.

The last section in this volume aims to addressprecisely these questions concerning the cognitiveunderpinnings of musical disorders. The section be-gins by exploring memory limitations in personswith congenital amusia, a topic introduced by DallaBella et al., who outline a model of vocal produc-tion and its impairment in amusia and suggest thatmemory can be a source of disorders in musicalproduction.121 Stewart, in collaboration with SusanAnderson et al., describe and present preliminaryresults from a musical intervention with five indi-viduals diagnosed with congenital amusia.122 Louiexplores further the notion of learning in musi-cal disabilities by reporting a novel investigation ofrapid statistical learning abilities and its disruptionin tone-deaf individuals.123 This statistical learningapproach is followed up in the last chapter by Peretzet al., who report five novel experiments showingthat persons with amusia can learn novel words aseasily as controls, whereas they systematically fail onmusical materials.124

Taken together, the aim of this section is to pro-vide a comprehensive review of musical disorders,bring novel results to light regarding learning andmemory and their interactions in normal and disor-dered brains, and ultimately to offer new perspec-tives toward the fundamental questions regardingneuroplasticity and the domain generality versusdomain specificity of music.

Conclusion

It is evident from this introduction that the field ofmusic neuroscience is rich and varied in ideas, ap-proaches, implications, and potential applications.Much of the work is interdisciplinary in nature, in-cluding strong links with psychology, neurology,and clinical practice, and growing links with mu-sic performance, education, and therapy. Althoughthe papers included here certainly reflect the currentstate of the art, it must also be emphasized that theyrepresent only a small fraction of ongoing researchinternationally and only hint at the research beingplanned and prepared. The future of our under-standing of the underlying neurobiology of humanmusical ability and behavior looks healthy, exciting,and promising, from the first milliseconds of audi-tory perception in infants to long-term training, in-dividual musical preferences, and cultural diversity.We look forward with anticipation to The Neuro-sciences and Music V, planned for France in 2014.

Conflicts of interest

The authors declare no conflicts of interest.

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