Cognitive neuroscience of music 1

Cognitive neuroscience of musicThe cognitive neuroscience of music is the scientific study of brain-based mechanisms involved in the cognitiveprocesses underlying music. These behaviours include music listening, performing, composing, reading, writing, andancillary activities. It also is increasingly concerned with the brain basis for musical aesthetics and musical emotion.Scientists working in this field may have training in cognitive neuroscience, neurology, neuroanatomy, psychology,music theory, computer science, and other allied fields.Cognitive neuroscience of music is distinguished from related fields such as music psychology, music cognition andcognitive musicology in its reliance on direct observations of the brain, using such techniques as functional magneticresonance imaging (fMRI), transcranial magnetic stimulation (TMS), magnetoencephalography (MEG),electroencephalography (EEG), and positron emission tomography (PET).

Neurological Bases

PitchFor each pitch there is a corresponding part of the tonotopically organized basilar membrane in the inner ear whichresponds to the sound and sends a signal to the auditory cortex. Sections of cells in the cortex are responsive tocertain frequencies, which range from very low to very high in pitches.[1] This organization may not be stable andthe specific cells that are responsive to different pitches may change over days or months.[2]

Melody Processing in the Secondary Auditory CortexStudies suggest that individuals are capable of automatically detecting a difference or anomaly in a melody such asan out of tune pitch which does not fit with their previous music experience. This automatic processing occurs in thesecondary auditory cortex. Brattico, Tervaniemi, Naatanen, and Peretz (2006) performed one such study todetermine if the detection of tones that do not fit an individual's expectations can occur automatically.[3] Theyrecorded event-related potentials (ERPs) in nonmusicians as they were presented unfamiliar melodies with either anout of tune pitch or an out of key pitch while participants were either distracted from the sounds or attending to themelody. Both conditions revealed an early frontal negativity independent of where attention was directed. Thisnegativity originated in the auditory cortex, more precisely in the supratemporal lobe (which corresponds with thesecondary auditory cortex) with greater activity from the right hemisphere. The negativity response was larger forpitch that was out of tune than that which was out of key. Ratings of musical incongruity were higher for out of tunepitch melodies than for out of key pitch. In the focused attention condition, out of key and out of tune pitchesproduced late parietal positivity. The findings of Brattico et al. (2006) suggest that there is automatic and rapidprocessing of melodic properties in the secondary auditory cortex.[3] The findings that pitch incongruities weredetected automatically, even in processing unfamiliar melodies, suggests that there is an automatic comparison ofincoming information with long term knowledge of musical scale properties, such as culturally influenced rules ofmusical properties (common chord progressions, scale patterms, etc.) and individual expectations of how the melodyshould proceed.

Cognitive neuroscience of music 2

Role of Right Auditory Cortex in Fine Pitch Resolution

The primary auditory cortex is one of the main areas associated withsuperior pitch resolution.

The right secondary auditory cortex has finer pitchresolution than the left. Hyde, Peretz and Zatorre(2008) used functional magnetic resonance imaging(fMRI) in their study to test the involvement of rightand left auditory cortical regions in frequencyprocessing of melodic sequences.[4] As well as findingsuperior pitch resolution in the right secondary auditorycortex, specific areas found to be involved were theplanum temporale (PT) in the secondary auditorycortex, and the primary auditory cortex in the medialsection of Heschl’s gyrus (HG).

Many neuroimaging studies have found evidence of theimportance of right secondary auditory regions inaspects of musical pitch processing, such as melody.[5] Many of these studies such as one by Patterson, Uppenkamp,Johnsrude and Griffiths (2002) also find evidence of a hierarchy of pitch processing. Patterson et al. (2002) usedspectrally matched sounds which produced: no pitch, fixed pitch or melody in an fMRI study and found that allconditions activated HG and PT. Sounds with pitch activated more of these regions than sounds without. When amelody was produced activation spread to the superior temporal gyrus (STG) and planum polare (PP). These resultssupport the existence of a pitch processing hierarchy.

RhythmThe belt and parabelt areas of the right hemisphere are involved in processing rhythm. When individuals arepreparing to tap out a rhythm of regular intervals (1:2 or 1:3) the left frontal cortex, left parietal cortex, and rightcerebellum are all activated. With more difficult rhythms such as a 1:2.5, more areas in the cerebral cortex andcerebellum are involved.[6] EEG recordings have also shown a relationship between brain electrical activity andrhythm perception. Snyder and Large (2005) performed a study examining rhythm perception in human subjects,finding that activity in the gamma band (20 – 60 Hz) corresponds to the 'beats' in a simple rhythm. Two types ofgamma activity were found by Snyder et al: (2005); induced gamma activity, and evoked gamma activity. Evokedgamma activity was found after the onset of each tone in the rhythm; this activity was found to be phase-locked(peaks and troughs were directly related to the exact onset of the tone) and did not appear when a gap (missed beat)was present in the rhythm. Induced gamma activity, which was not found to be phase-locked, was also found tocorrespond with each beat. However, induced gamma activity did not subside when a gap was present in the rhythm,indicating that induced gamma activity may possibly serve as a sort of internal metronome independent of auditoryinput.

Cognitive neuroscience of music 3

TonalityThe right auditory cortex is primary involved in perceiving pitch, and parts of harmony, melody and rhythm.[6] Onestudy by Peter Janata found that there are tonally sensitive areas in the medial prefrontal cortex, the cerebellum, thesuperior Temporal sulci of both hemispheres and the Superior Temporal gyri (which has a skew towards the righthemisphere).

Music and LanguageCertain aspects of language and melody have been shown to be processed in near identical functional brain areas.Brown, Martinez and Parsons (2006) examined the neurological structural similarities between music andlanguage.[7] Utilizing positron emission tomography (PET), the findings showed that both linguistic and melodicphrases produced activation in almost identical functional brain areas. These areas included the primary motorcortex, supplementary motor area, Broca’s area, anterior insula, primary and secondary auditory cortices, temporalpole, basal ganglia, ventral thalamus and posterior cerebellum. Differences were found in lateralization tendencies aslanguage tasks favoured the left hemisphere, but the majority of activations were bilateral which producedsignificant overlap across modalities.[7]

Syntactical information mechanisms in both music and language have been shown to be processed similarly in thebrain. Jentschke, Koelsch, Sallat and Friederici (2008) conducted a study investigating the processing of music inchildren with specific language impairments (SLI).[8] Children with typical language development (TLD) showedERP patterns different than that of children with SLI which reflected their challenges to process music-syntacticregularities. Strong correlations between the ERAN amplitude and linguistic and musical abilities provide additionalevidence for the relationship of syntactical processing in music and language.[8]

However, production of melody and production of speech may be subserved by different neural networks. Stewart,Walsh, Frith and Rothwell (2001) studied the differences between speech production and song production usingtranscranial magnetic stimulation (TMS).[9] Stewart et al. found that TMS applied to the left frontal lobe disturbsspeech but not melody supporting the idea that they are subserved by different areas of the brain. The authorssuggest that a reason for the difference is that speech generation can be localized well but the underlyingmechanisms of melodic production cannot. Alternatively, it was also suggested that speech production may be lessrobust than melodic production and thus more susceptible to interference.[9]

Language processing is a function more of the left side of the brain than the right side, particularly Broca's Area andWernicke's area, though the roles played by the two sides of the brain in processing different aspects of language arestill unclear. Music is also processed by both the left and the right sides of the brain.[10][11] Recent evidence furthersuggest shared processing between language and music at the conceptual level.[12] It has also been found that, amongmusic conservatory students, the prevalence of absolute pitch is much higher for speakers of tone language, evencontrolling for ethnic background, showing that language influences how musical tones are perceived. [13][14]

Cognitive neuroscience of music 4

Musician vs. Non-musician processing

Professional piano players show less corticalactivation for complex finger movement tasks

due to structural differences in the brain.

Differences

Brain structure within musicians and non-musicians is distinctlydifferent. Gaser and Schlaug (2003) compared brain structures ofprofessional musicians with non-musicians and discovered gray mattervolume differences in motor, auditory and visual-spatial brainregions.[15] Specifically, positive correlations were discovered betweenmusician status (professional, amateur and non-musician) and graymatter volume in the primary motor and somatosensory areas,premotor areas, anterior superior parietal areas and in the inferiortemporal gyrus bilaterally. This strong association between musicianstatus and gray matter differences supports the notion that musicians’brains show use-dependent structural changes.[16] Due to the distinctdifferences in several brain regions, it is unlikely that these differencesare innate but rather due to the long-term acquisition and repetitiverehearsal of musical skills.

Brains of musicians also show functional differences from those ofnon-musicians. Krings, Topper, Foltys, Erberich, Sparing, Willmes andThron (2000) utilized fMRI to study brain area involvement ofprofessional piano players and a control group while performingcomplex finger movements.[17] Krings et al. found that the professional piano players showed lower levels of corticalactivation in motor areas of the brain. It was concluded that a lesser amount of neurons needed to be activated for thepiano players due to long-term motor practice which results in the different cortical activation patterns. Koeneke,Lutz, Wustenberg and Jancke (2004) reported similar findings in keyboard players.[18] Skilled keyboard players anda control group performed complex tasks involving unimanual and bimanual finger movements. During taskconditions, strong hemodynamic responses in the cerebellum were shown by both non-musicians and keyboardplayers, but non-musicians showed the stronger response. This finding indicates that different cortical activationpatterns emerge from long-term motor practice. This evidence supports previous data showing that musicians requirefewer neurons to perform the same movements.

Musicians have been shown to have significantly more developed left planum temporales, and have also shown tohave a greater word memory (Chan et al.). Chan’s study controlled for age, grade point average and years ofeducation and found that when given a 16 word memory test, the musicians averaged one to two more words abovetheir non musical counterparts.

SimilaritiesStudies have shown that the human brain has an implicit musical ability.[19][20] Koelsch, Gunter, Friederici and Schoger (2000) investigated the influence of preceding musical context, task relevance of unexpected chords and the degree of probability of violation on music processing in both musicians and non-musicians.[19] Findings showed that the human brain unintentionally extrapolates expectations about impending auditory input. Even in non-musicians, the extrapolated expectations are consistent with music theory. The ability to process information musically supports the idea of an implicit musical ability in the human brain. In a follow-up study, Koelsch, Schroger, and Gunter (2002) investigated whether ERAN and N5 could be evoked preattentively in non-musicians.[20] Findings showed that both ERAN and N5 can be elicited even in a situation where the musical stimulus is ignored by the listener indicating that there is a highly differentiated preattentive musicality in the human

Cognitive neuroscience of music 5

brain.

Gender DifferencesMinor neurological differences with regard to hemispheric processing exist between brains of males and females.Koelsch, Maess, Grossmann and Friederici (2003) investigated music processing through EEG and ERPs anddiscovered gender differences.[21] Findings showed that females process music information bilaterally and malesprocess music with a right-hemispheric predominance. However, the early negativity of males was also present overthe left hemisphere. This indicates that males do not exclusively utilize the right hemisphere for musical informationprocessing. In a follow-up study, Koelsch, Grossman, Gunter, Hahne, Schroger and Friederici (2003) found that boysshow lateralization of the early anterior negativity in the left hemisphere but found a bilateral effect in girls.[22] Thisindicates a developmental effect as early negativity is lateralized in the right hemisphere in men and in the lefthemisphere in boys.

Handedness DifferencesIt has been found that subjects who are lefthanded, particularly those who are also ambidextrous, perform better thanrighthanders on short term memory for the pitch.[23][24] It was hypothesized that this handedness advantage is due tothe fact that lefthanders have more duplication of storage in the two hemispheres than do righthanders. Other workhas shown that there are pronounced differences between righthanders and lefthanders (on a statistical basis) in howmusical patterns are perceived, when sounds come from different regions of space. This has been found, forexample, in the Octave illusion [25][26] and the Scale illusion.[27][28]

Musical ImageryMusical imagery refers to the experience of replaying music by imagining it inside the head.[29] Musicians show asuperior ability for musical imagery due to intense musical training.[30] Herholz, Lappe, Knief and Pantev (2008)investigated the differences in neural processing of a musical imagery task in musicians and non-musicians. Utilizingmagnetoencephalography (MEG), Herholz et al. examined differences in the processing of a musical imagery taskwith familiar melodies in musicians and non-musicians. Specifically, the study examined whether the mismatchnegativity (MMN) can be based solely on imagery of sounds. The task involved participants listening to thebeginning of a melody, continuation of the melody in his/her head and finally hearing a correct/incorrect tone asfurther continuation of the melody. The imagery of these melodies was strong enough to obtain an early preattentivebrain response to unanticipated violations of the imagined melodies in the musicians. These results indicate similarneural correlates are relied upon for trained musicians imagery and perception. Additionally, the findings suggestthat modification of the imagery mismatch negativity (iMMN) through intense musical training results inachievement of a superior ability for imagery and preattentive processing of music.Perceptual musical processes and musical imagery may share a neural substrate in the brain. A PET study conductedby Zatorre, Halpern, Perry, Meyer and Evans (1996) investigated cerebral blood flow (CBF) changes related toauditory imagery and perceptual tasks.[31] These tasks examined the involvement of particular anatomical regions aswell as functional commonalities between perceptual processes and imagery. Similar patterns of CBF changesprovided evidence supporting the notion that imagery processes share a substantial neural substrate with relatedperceptual processes. Bilateral neural activity in the secondary auditory cortex was associated with both perceivingand imagining songs. This implies that within the secondary auditory cortex, processes underlie thephenomenological impression of imagined sounds. The supplementary motor area (SMA) was active in both imageryand perceptual tasks suggesting covert vocalization as an element of musical imagery. CBF increases in the inferiorfrontal polar cortex and right thalamus suggest that these regions may be related to retrieval and/or generation ofauditory information from memory.

Cognitive neuroscience of music 6

Absolute Pitch

Musicians possessing absolute pitch canidentify the pitch of musical tones

without external reference.

Absolute pitch (AP) is defined as the ability to identify the pitch of a musicaltone or to produce a musical tone at a given pitch without the use of anexternal reference pitch.[32] Neuroscientific research has not discovered adistinct activation pattern common for possessors of AP. Zatorre, Perry,Beckett, Westbury and Evans (1998) examined the neural foundations of APusing functional and structural brain imaging techniques.[33] Positronemission tomography (PET) was utilized to measure cerebral blood flow(CBF) in musicians possessing AP and musicians lacking AP. Whenpresented with musical tones, similar patterns of increased CBF in auditorycortical areas emerged in both groups. AP possessors and non-AP subjects demonstrated similar patterns of leftdorsolateral frontal activity when they performed relative pitch judgments. However, in non-AP subjects activationin the right inferior frontal cortex was present whereas AP possessors showed no such activity. This finding suggeststhat musicians with AP do not need access to working memory devices for such tasks. These findings imply thatthere is no specific regional activation pattern unique to AP. Rather, the availability of specific processingmechanisms and task demands determine the recruited neural areas.

EmotionEmotions induced by music activate similar frontal brain regions compared to emotions elicited by other stimuli.[34]

Schmidt and Trainor (2001) discovered that valence (i.e. positive vs. negative) of musical segments wasdistinguished by patterns of frontal EEG activity.[35] Joyful and happy musical segments were associated withincreases in left frontal EEG activity whereas fearful and sad musical segments were associated with increases inright frontal EEG activity. Additionally, the intensity of emotions was differentiated by the pattern of overall frontalEEG activity. Overall frontal region activity increased as affective musical stimuli became more intense.[35]

Music is able to create an incredibly pleasurable experience that can be described as “chills”.[36] Blood and Zatorre(2001) used PET to measure changes in cerebral blood flow while participants listened to music that they knew togive them the “chills” or any sort of intensely pleasant emotional response. They found that as these chills increase,many changes in cerebral blood flow are seen in brain regions such as the amygdala, orbitofrontal cortex, ventralstriatum, midbrain, and the ventral medial prefrontal cortex. Many of these areas appear to be linked to reward andmotivation, emotion and arousal and are also activated in other pleasurable situations.[36] Nucleus accumbens (a partof striatum) is involved in both music related emotions, as well as rhythmic timing.When unpleasant melodies are played, the posterior cingulate cortex activates, which indicates a sense of conflict oremotional pain.[6] The right hemisphere has also been found to be correlated with emotion, which can also activateareas in the cingluate in times of emotional pain, specifically social rejection (Eisenberger). This evidence, alongwith observations, has led many musical theorists, philosophers and neuroscientists to link emotion with tonality.This seems almost obvious because the tones in music seem like a characterization of the tones in human speech,which indicate emotional content. The vowels in the phonemes of a song are elongated for a dramatic effect, and itseems as though musical tones are simply exaggerations of the normal verbal tonality.

Cognitive neuroscience of music 7

Memory

Neuropsychology of Musical MemoryMusical memory involves both explicit and implicit memory systems.[37] Explicit musical memory is furtherdifferentiated between episodic (where, when and what of the musical experience) and semantic (memory for musicknowledge including facts and emotional concepts). Implicit memory centers on the ‘how’ of music and involvesautomatic processes such as procedural memory and motor skill learning – in other words skills critical for playingan instrument. Samson and Baird (2009) found that the ability of musicians with Alzheimer’s Disease to play aninstrument (implicit procedural memory) may be preserved.

Neural correlates of Musical MemoryA PET study looking into the neural correlates of musical semantic and episodic memory found distinct activationpatterns.[38] Semantic musical memory involves the sense of familiarity of songs. The semantic memory for musiccondition resulted in bilateral activation in the medial and orbital frontal cortex, as well as activation in the leftangular gyrus and the left anterior region of the middle temporal gyri. These patterns support the functionalasymmetry favouring the left hemisphere for semantic memory. Left anterior temporal and inferior frontal regionsthat were activated in the musical semantic memory task produced activation peaks specifically during thepresentation of musical material, suggestion that these regions are somewhat functionally specialized for musicalsemantic representations.Episodic memory of musical information involves the ability to recall the former context associated with a musicalexcerpt.[38] In the condition invoking episodic memory for music, activations were found bilaterally in the middleand superior frontal gyri and precuneus, with activation predominant in the right hemisphere. Other studies havefound the precuneus to become activated in successful episodic recall.[39] As it was activated in the familiar memorycondition of episodic memory, this activation may be explained by the successful recall of the melody.When it comes to memory for pitch, there appears to be a dynamic and distributed brain network subserves pitchmemory processes. Gaab, Gaser, Zaehle, Jancke and Schlaug (2003) examined the functional anatomy of pitchmemory using functional magnetic resonance imaging (fMRI).[40] An analysis of performance scores in a pitchmemory task resulted in a significant correlation between good task performance and the supramarginal gyrus(SMG) as well as the dorsolateral cerebellum. Findings indicate that the dorsolateral cerebellum may act as a pitchdiscrimination processor and the SMG may act as a short-term pitch information storage site. The left hemispherewas found to be more prominent in the pitch memory task than the right hemispheric regions.

Therapeutic effects of music on memoryMusical training has been shown to aid memory. Altenmuller et al. studied the difference between active and passivemusical instruction and found both that over a longer (but not short) period of time, the actively taught studentsretained much more information than the passively taught students. The actively taught students were also found tohave greater cerebral cortex activation. It should also be noted that the passively taught students weren’t wastingtheir time; they, along with the active group, displayed greater left hemisphere activity, which is typical in trainedmusicians.[41]

Cognitive neuroscience of music 8

DevelopmentThe musical four year olds have been found to have compared to one greater left hemisphere intrahemisphericcoherence.[41] Musicians have been found to have more developed anterior portions of the corpus callosum in astudy by Cowell et al. in 1992 . This was confirmed by a study by Schlaug et al. in 1995 who found that classicalmusicians between the ages of 21 and 36 have significantly greater anterior corpora callosa than the non-musicalcontrol. Schlaug also found that there was a strong correlation of musical exposure before the age of seven, and agreat increase in the size of the corpus callosum.[41] These fibers join together the left and right hemispheres andindicate an increased relaying between both sides of the brain. This suggests the merging between the spatial-emotiono-tonal processing of the right brains and the linguistical processing of the left brain. This large relayingacross many different areas of the brain might contribute to music’s ability to aid in memory function.

Impairment

Focal Hand DystoniaFocal hand dystonia is a task-related movement disorder associated with occupational activities that requirerepetitive hand movements.[42] Focal hand dystonia is associated with abnormal processing in the premotor andprimary sensorimotor cortices. An fMRI study examined five guitarists with focal hand dystonia.[43] The studyreproduced task-specific hand dystonia by having guitarists use a real guitar neck inside the scanner as well asperforming a guitar exercise to trigger abnormal hand movement. The dystonic guitarists showed significantly moreactivation of the contralateral primary sensorimotor cortex as well as a bilateral underactivation of premotor areas.This activation pattern represents abnormal recruitment of the cortical areas involved in motor control. Even inprofessional musicians, widespread bilateral cortical region involvement is necessary to produce complex handmovements such as scales and arpeggios. The abnormal shift from premotor to primary sensorimotor activationdirectly correlates with guitar-induced hand dystonia.

Music AgnosiaMusic agnosia, an auditory agnosia, is a syndrome of selective impairment in music recognition.[44] Three cases ofmusic agnosia are examined by Dalla Bella and Peretz (1999); C.N., G.L., and I.R.. All three of these patientssuffered bilateral damage to the auditory cortex which resulted in musical difficulties while speech understandingremained intact. Their impairment is specific to the recognition of once familiar melodies. They are spared inrecognizing environmental sounds and in recognizing lyrics. Peretz (1996) has studied C.N.’s music agnosia furtherand reports an initial impairment of pitch processing and spared temporal processing.[45] C.N. later recovered inpitch processing abilities but remained impaired in tune recognition and familiarity judgments.Musical agnosias may be categorized based on the process which is impaired in the individual.[46] Apperceptivemusic agnosia involves an impairment at the level of perceptual analysis involving an inability to encode musicalinformation correctly. Associative music agnosia reflects an impaired representational system which disrupts musicrecognition. Many of the cases of music agnosia have resulted from surgery involving the middle cerebral artery.Patient studies have surmounted a large amount of evidence demonstrating that the left side of the brain is moresuitable for holding long-term memory representations of music and that the right side is important for controllingaccess to these representations. Associative music agnosias tend to be produced by damage to the left hemisphere,while apperceptive music agnosia reflects damage the to right hemisphere.

Cognitive neuroscience of music 9

Congenital AmusiaCongenital amusia, otherwise known as tone deafness, is a term for lifelong musical problems which are notattributable to mental retardation, lack of exposure to music or deafness, or brain damage after birth .[47] Amusicbrains have been found in fMRI studies to have less white matter and thicker cortex than controls in the right inferiorfrontal cortex. These differences suggest abnormal neuronal development in the auditory cortex and inferior frontalgyrus, two areas which are important in musical-pitch processing.Studies on those with amusia suggest different processes are involved in speech tonality and musical tonality.Congenital amusics lack the ability to distinguish between pitches and so are for example unmoved by dissonanceand playing the wrong key on a piano. They also cannot be taught to remember a melody or to recite a song;however, they are still capable of hearing the intonation of speech, for example, distinguishing between “You speakFrench” and “You speak French?” when spoken.

Amygdala damage

Damage to the amygdala may impair recognition ofscary music.

Damage to the amygdala has selective emotional impairments onmusical recognition. Gosselin, Peretz, Johnsen and Adolphs(2007) studied S.M., a patient with bilateral damage of theamygdala with the rest of the temporal lobe undamaged and foundthat S.M. was impaired in recognition of scary and sad music.[48]

S.M.’s perception of happy music was normal, as was her ability touse cues such as tempo to distinguish between happy and sadmusic. It appears that damage specific to the amygdala canselectively impair recognition of scary music.

Selective Deficit in Music Reading

Specific musical impairments may result from brain damageleaving other musical abilities intact. Cappelletti, Waley-Cohen,Butterworth and Kopelman (2000) studied a single case study ofpatient P.K.C., a professional musician who sustainted damage tothe left posterior temporal lobe as well as a small rightoccipitotemporal lesion.[49] After sustaining damage to theseregions, P.K.C. was selectively impaired in the areas of reading, writing and understanding musical notation butmaintained other musical skills. The ability to read aloud letters, words, numbers and symbols (including musicalones) was retained. However, P.K.C. was unable to read aloud musical notes on the staff regardless of whether thetask involved naming with the conventional letter or by singing or playing. Yet despite this specific deficit, P.K.C.retained the ability to remember and play familiar and new melodies.

Auditory ArrhythmiaArrhythmia in the auditory modality is defined as a disturbance of rhythmic sense; and includes deficits such as theinability to rhythmically perform music, the inability to keep time to music and the inability to discriminate betweenor reproduce rhythmic patterns.[50] A study investigating the elements of rhythmic function examined Patient H.J.,who acquired arrhythmia after sustaining a right temporoparietal infarct.[50] Damage to this region impaired H.J.’scentral timing system which is essentially the basis of his global rhythmic impairment. H.J. was unable to generatesteady pulses in a tapping task. These findings suggest that keeping a musical beat relies on functioning in the righttemporal auditory cortex.

Cognitive neuroscience of music 10

References[1] Arlinger, S.; Elberling, C.; Bak, C.; Kofoed, B.; Lebech, J.; Saermark, K. (1982). "Cortical magnetic fields evoked by frequency glides of a

continuous tone". EEG & Clinical Neurophysiology 54 (6): 642–653. doi:10.1016/0013-4694(82)90118-3.[2] Janata P, Birk J, Van Horn J, Leman M, Tillmann B, Bharucha J. (2002). "The cortical topography of tonal structures underlying Western

music". Science 298 (5601): 2167–70. doi:10.1126/science.1076262. PMID 12481131.[3] Brattico, E.; Tervaniemi, M.; Naatanen, R.; Peretz, I. (2006). "Musical scale properties are automatically processed in the human auditory

cortex". Brain Research 1117 (1): 162–174. doi:10.1016/j.brainres.2006.08.023. PMID 16963000.[4] Hyde, K. L; Peretz, I.; Zatorre, R. J. (2008). "Evidence for the role of the right auditory cortex in fine pitch resolution". Neuropsychologia 46

(2): 632–639. doi:10.1016/j.neuropsychologia.2007.09.004. PMID 17959204.[5] Patterson, R. D.; Uppenkamp, S.; Johnsrude, I. S.; Griffiths, T. D. (2002). "The processing of temporal pitch and melody information in

auditory cortex". Neuron 36 (4): 767–776. doi:10.1016/S0896-6273(02)01060-7. PMID 12441063.[6] Tramo MJ. (2001). "Biology and music. Music of the hemispheres". Science 291 (5501): 54–6.

doi:10.1126/science.10.1126/SCIENCE.1056899. PMID 11192009.[7] Brown, S.; Martinez, M.J.; Parsons, L.M. (2006). "Music and language side by side in the brain: a PET study of the generation of melodies

and sentences". European Journal of Neuroscience 23 (10): 2791–2803. doi:10.1111/j.1460-9568.2006.04785.x. PMID 16817882.[8] Jentschke, S.; Koelsch, S.; Sallat, S.; Friederici, A.D. (2008). "Children with specific language impairment also show impairment of

music-syntactic processing". Journal of Cognitive Neuroscience 20 (11): 1940–1951. doi:10.1162/jocn.2008.20135. PMID 18416683.[9] Stewart, L.; Walsh, V.; Frith, U.; Rothwell, J. (2001). "Transcranial magnetic stimulation produces speech arrest but not song arrest". Annals

New York Academy of Sciences 930: 433–35. doi:10.1111/j.1749-6632.2001.tb05762.x.[10] Koelsch, S., Gunter, T., Cramon, D., Zysset, S., Lohmann, G. & Friederici, A. (2002). "Bach Speaks: A Cortical Language-Network Serves

the Processing of Music". NeuroImage 17 (2): 956–966. doi:10.1006/nimg.2002.1154. PMID 12377169.[11] Brown, S., Martinez, M. & Parsons, L. (2006). "Music and language side by side in the brain: a PET study of the generation of melodies and

sentences". European Journal of Neuroscience 23 (10): 2791–2803. doi:10.1111/j.1460-9568.2006.04785.x. PMID 16817882.[12] Daltrozzo, J., Schön, D. (2009). Conceptual processing in music as revealed by N400 effects on words and musical targets. Journal of

Cognitive Neuroscience, 21(10): 1882-1892. (http:/ / daltrozzo. net78. net/ papers/ Daltrozzo_Schon_2009a. pdf)[13] Deutsch, D., Henthorn, T., Marvin, E., & Xu H-S (2006). "Absolute pitch among American and Chinese conservatory students: Prevalence

differences, and evidence for a speech-related critical period". Journal of the Acoustical Society of America 119 (2): 719–722.doi:10.1121/1.2151799. PMID 16521731. PDF Document (http:/ / philomel. com/ pdf/ JASA-2006_119_719-722. pdf)

[14] Deutsch, D., Dooley, K., Henthorn, T. and Head, B. (2009). "Absolute pitch among students in an American music conservatory:Association with tone language fluency". Journal of the Acoustical Society of America 125 (4): 2398–2403. doi:10.1121/1.3081389.PMID 19354413. Weblink (http:/ / scitation. aip. org/ vsearch/ servlet/ VerityServlet?KEY=ASADL& smode=strresults& sort=rel&maxdisp=25& threshold=0& pjournals=ARLOFJ,JASMAN,NOCOAN,SOUCAU,PMARCW,ATCODK,ASASTR& possible1=Absolute+pitch+ among+ students+ in+ an+ American+ music+ conservatory& possible1zone=article& OUTLOG=NO& viewabs=JASMAN&key=DISPLAY& docID=2& page=0& chapter=0) PDF Document (http:/ / philomel. com/ pdf/ JASA-2009_125_2398-2403. pdf)

[15] Gaser, C.; Schlaug, G. (2003). "Brain structures differ between musicians and non-musicians". The Journal of Neuroscience 23 (27):9240–9245. PMID 14534258.

[16] Croom, A.M. (2012). "Music, neuroscience, and the psychology of wellbeing: a précis". Frontiers in Theoretical and PhilosophicalPsychology 2 (393): 1-15. doi:10.3389/fpsyg.2011.00393.

[17] Krings, T.; Topper, R.; Foltys, H.; Erberich, S.; Sparing, R.; Willmes, K.; Thron, A. (1999). "Cortical activation patterns during complexmotor tasks in piano players and control subjects. A functional magnetic resonance imaging study". Neuroscience Letters 278 (3): 189–193.doi:10.1016/S0304-3940(99)00930-1. PMID 10653025.

[18] Koeneke, S.; Lutz, K.; Wustenberg, T.; Jancke, L. (2004). "Long-term training affects cerebellar processing in skilled keyboard players".Neuroreport 15 (8): 1279–1282. doi:10.1097/01.wnr.0000127463.10147.e7. PMID 15167549.

[19] Koelsch, S.; Gunter, T.; Friederici, A.D.; Schoger, E. (2000). "Brain indices of music processing: "nonmusicians" are musical". Journal ofCognitive Neuroscience 12 (3): 520–541. doi:10.1162/089892900562183. PMID 10931776.

[20] Koelsch, S.; Schroger, E.; Gunter, T. (2002). "Music matters: preattentive musicality of the human brain". Psychophysiology 39 (1): 38–48.doi:10.1111/1469-8986.3910038. PMID 12206294.

[21] Koelsch, S.; Maess, B.; Grossmann, T.; Friederici, A.D. (2003). "Electric brain responses reveal gender differences in music processing".Neuroreport 14 (5): 709–713. doi:10.1097/00001756-200304150-00010. PMID 12692468.

[22] Koelsch, S.; Grossmann, T.; Gunter, T.C.; Hahne, A.; Schroger, E.; Friederici, A.D. (2003). "Children processing music: electric brainresponses reveal musical competence and gender differences". Journal of Cognitive Neuroscience 15 (5): 683–693.doi:10.1162/jocn.2003.15.5.683. PMID 12965042.

[23] Deutsch, D. (1978). "Pitch memory: An advantage for the lefthanded.". Science 199 (4328): 559–560. doi:10.1126/science.622558.PMID 622558. PDF Document (http:/ / philomel. com/ pdf/ Science-1978_199_559-560. pdf)

[24] Deutsch, D. (1980). "Handedness and Memory for Tonal Pitch. In J. Herron (Ed.)". Neuropsychology of Lefthandedness: 263–271. PDFDocument (http:/ / philomel. com/ pdf/ Ch-Handedness-1980. pdf)

[25] Deutsch, D. (1974). "An auditory illusion". Nature 251 (5473): 307–309. doi:10.1038/251307a0. PMID 4427654. Weblink (http:/ / www.nature. com/ nature/ journal/ v251/ n5473/ abs/ 251307a0. html) PDF Document (http:/ / philomel. com/ pdf/ Nature-1974_251_307-309. pdf)

Cognitive neuroscience of music 11

[26] Deutsch, D. (1983). "The octave illusion in relation to handedness and familial handedness background.". Neuropsychologia 21 (3):289–293. doi:10.1016/0028-3932(83)90047-7. PMID 6877583. PDF Document (http:/ / philomel. com/ pdf/ Neuropsych-1983_21_289-293.pdf)

[27] Deutsch, D. (1975). "Two-channel listening to musical scales". Journal of the Acoustical Society of America 57 (5): 1156–1160.doi:10.1121/1.380573. PMID 1127169. Weblink (http:/ / scitation. aip. org/ vsearch/ servlet/ VerityServlet?KEY=ASADL&smode=strresults& sort=chron& maxdisp=25& threshold=0& pjournals=journals& pjournals=JASMAN& pjournals=ARLOFJ&pjournals=NOCOAN& pjournals=SOUCAU& possible1=Two-channel listening to musical scales. & possible1zone=title& OUTLOG=NO&viewabs=JASMAN& key=DISPLAY& docID=1& page=1& chapter=0) PDF Document (http:/ / philomel. com/ pdf/JASA-1975_57_1156-1160. pdf)

[28] Deutsch, D. (1999). "Grouping mechanisms in music. In D. Deutsch (Ed.)". The psychology of music, 2nd Edition: 299–348. PDF Document(http:/ / philomel. com/ pdf/ PsychMus_Ch9. pdf)

[29] Halpern, A.R (2001). "Cerebral substrates of musical imagery". Annals New York Academy of Sciences 930: 179–192.doi:10.1111/j.1749-6632.2001.tb05733.x.

[30] Herholz, S.C.; Lappe, C.; Knief, A.; Pantev, C. (2008). "Neural basis of music imagery and the effect of musical expertise". EuropeanJournal of Neuroscience 28 (11): 2352–2360. doi:10.1111/j.1460-9568.2008.06515.x. PMID 19046375.

[31] Zatorre, R.J.; Halpern, A.R.; Perry, D.W.; Meyer, E.; Evans, A.C. (1996). "Hearing in the mind's ear: A PET investigation of musicalimagery and perception". Journal of Cognitive Neuroscience 8 (1): 29–46. doi:10.1162/jocn.1996.8.1.29.

[32] Takeuchi, A.H.; Hulse, S.H. (1993). "Absolute pitch". Psychological Bulletin 113 (2): 345–361. doi:10.1037/0033-2909.113.2.345.PMID 8451339.

[33] Zatorre, R.J.; Perry, D.W.; Beckett, C.A.; Westbury, C.F.; Evans, A.C. (1998). "Functional anatomy of musical processing in listeners withabsolute pitch and relative pitch". Neurobiology 95: 3172–3177.

[34] Croom, A.M. (2012). "Music, neuroscience, and the psychology of wellbeing: a précis". Frontiers in Theoretical and PhilosophicalPsychology 2 (393): 1-15. doi:10.3389/fpsyg.2011.00393.

[35] Schmidt, L.A.; Trainor, L.J. (2001). "Frontal brain electrical activity (EEG) distinguishes valence and intensity of musical emotions".Cognition and Emotion 15 (4): 487–500.

[36] Blood, A. J.; Zatorre, R. J. (2001). "Intensely pleasurable responses to music correlate with activity in brain regions implicated in reward andemotion". Proceedings of the National Academy of Sciences of the United States of America 98 (20): 11818–11823.doi:10.1073/pnas.191355898. PMC 58814. PMID 11573015.

[37] Baird, A.; Samson, S. (2009). "Memory for music in Alzheimer's Disease: Unforgettable?.". Neuropsychological Review 19: 85–101.doi:10.1007/s11065-009-9085-2.

[38] Platel, H.; Baron, J-C.; Desgranges, B.; Bernard, F.; Eustache, F. (2003). "Semantic and episodic memory of music are subserved by distinctneural networks". NeuroImage 20 (1): 244–256. doi:10.1016/S1053-8119(03)00287-8. PMID 14527585.

[39] Kapur, S.; Craik, F. I. M.; Jones, C.; Brown, G. M.; Houle, S.; Tulving, E. (1995). "Functional role of the prefrontal cortex in retrieval ofmemories: A PET study". NeuroReport 6 (14): 1880–1884. doi:10.1097/00001756-199510020-00014. PMID 8547589.

[40] Gaab, N; Gaser, C.; Zaehle, T.; Jancke, L.; Schlaug, G. (2003). "Functional anatomy of pitch memory – and fMRI study with sparsetemporal sampling". Neuroimage 19 (4): 1417–1426. doi:10.1016/S1053-8119(03)00224-6. PMID 12948699.

[41] Strickland JS. (2001). "Music and the Brain in Childhood Development" (http:/ / findarticles. com/ p/ articles/ mi_qa3614/ is_200101/ai_n8950759/ ). Childhood Education 78: 100–3. .

[42] Chen, R.; Hallett, M. (1998). "Focal dystonia and repetitive motion disorders". Occupational health and industrial medicine 39 (3): 122.[43] Pujol, J.; Roset-Llobet, J.; Rosines-Cubells, D.; Deus, J.; Narberhaus, B.; Valls-Sole, J.; Capdevila, A.; Pascual-Leone, A. (2000). "Brain

cortical activation during guitar-induced hand dystonia studied by functional MRI". Neuroimage 12 (3): 257–267.doi:10.1006/nimg.2000.0615. PMID 10944408.

[44] Dalla Bella, S.; Peretz, I. (1999). "Music agnosias: Selective impairments of music recognition after brain damage". Journal of New MusicResearch 28 (3): 209–216. doi:10.1076/jnmr.28.3.209.3108.

[45] Peretz, I. (1996). "Can we lose memory for music? A case of music agnosia in a nonmusician". Journal of Cognitive Neuroscience 8 (6):481–496. doi:10.1162/jocn.1996.8.6.481.

[46] Ayotte, J.; Peretz, I.; Rousseau, I.; Bard, C.; Bojanowski, M. (2000). "Patterns of music agnosia associated with middle cerebral arteryinfarcts". Brain 123: 1926–1938. doi:10.1093/brain/123.9.1926. PMID 10960056.

[47] Peretz, I. (2008). "Musical disorders". Current Directions in Psychological Science 17 (5): 329–333.doi:10.1111/j.1467-8721.2008.00600.x.

[48] Gosselin, N.; Peretz, I.; Johnsen, E.; Adolphs, R. (2007). "Amygdala damage impairs emotion recognition from music". Neuropsychologia45 (2): 236–244. doi:10.1016/j.neuropsychologia.2006.07.012. PMID 16970965.

[49] Cappelletti, M.; Waley-Cohen, H.; Butterworth, B.; Kopelman, M. (2000). "A selection loss of the ability to read and write music".Neurocase 6 (4): 321–332. doi:10.1080/13554790008402780.

[50] Wilson, S.J.; Pressing, J.L.; Wales, R.J. (2002). "Modelling rhythmic function in a musician post-stroke". Neuropsychologia 40 (8):1494–1505. doi:10.1016/S0028-3932(01)00198-1. PMID 11931954.

Cognitive neuroscience of music 12

Further reading• The neurosciences and music II : from perception to performance. New York, N.Y: New York Academy of

Sciences. 2005. ISBN 1573316105.

External links• Croom, Adam M. (2012). "Music, Neuroscience, and the Psychology of Wellbeing: A Précis." Frontiers in

Theoretical and Philosophical Psychology, 2, 393, 1-15 (http:/ / www. frontiersin. org/theoretical_and_philosophical_psychology/ 10. 3389/ fpsyg. 2011. 00393/ abstract).

• Deutsch, Diana (July 29, 2010). "Speaking in Tones: Music and Language Partner in the Brain" (http:/ / www.scientificamerican. com/ article. cfm?id=speaking-in-tones-jul10). Scientific American Mind.

• Weinberger, Norman M. (October 25, 2004). "Music and the Brain" (http:/ / www. sciam. com/ article.cfm?chanID=sa006& articleID=0007D716-71A1-1179-AF8683414B7F0000). Scientific American.

• Connection Science (2009). Special Issue "Music, Brain, & Cognition" 21(2-3). (http:/ / prod. informaworld. com/smpp/ title~db=all~content=g911586331)

Article Sources and Contributors 13

Article Sources and ContributorsCognitive neuroscience of music  Source: http://en.wikipedia.org/w/index.php?oldid=470527706  Contributors: Adamcroom, Aeternus, Bearcat, CharlesGillingham, David Eppstein, Dolfrog,Kimchi.sg, LilHelpa, Morton Shumway, Psyc4600, Rettetast, Rjwilmsi, Sj, Smallman12q, SolfegeNut, Thenthorn, WereSpielChequers, Woohookitty, 52 anonymous edits

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