semantic and episodic memory of music are subserved by distinct

Semantic and episodic memory of music are subservedby distinct neural networks

HervePlatel,a,* Jean-Claude Baron,b Beatrice Desgranges,a

Frederic Bernard,a and Francis Eustachea,c

a Inserm E.0218-Universite de Caen, Gip Cyceron, Laboratoire de Neuropsychologie, CHU Cote de Nacre, Caen, Franceb Department of Neurology, University of Cambridge, Cambridge UK

c Ecole Pratique des Hautes Etudes, CNRS 8581, Universite Rene Descartes, Paris 5, France

Received 26 September 2002; revised 14 February 2003; accepted 16 May 2003

Abstract

Numerous functional imaging studies have shown that retrieval from semantic and episodic memory is subserved by distinct neuralnetworks. However, these results were essentially obtained with verbal and visuospatial material. The aim of this work was to determinethe neural substrates underlying the semantic and episodic components of music using familiar and nonfamiliar melodic tunes. To studymusical semantic memory, we designed a task in which the instruction was to judge whether or not the musical extract was felt as “familiar.”To study musical episodic memory, we constructed two delayed recognition tasks, one containing only familiar and the other onlynonfamiliar items. For each recognition task, half of the extracts (targets) were presented in the prior semantic task. The episodic andsemantic tasks were to be contrasted by a comparison to two perceptive control tasks and to one another. Cerebral blood flow was assessedby means of the oxygen-15-labeled water injection method, using high-resolution PET. Distinct patterns of activations were found. First,regarding the episodic memory condition, bilateral activations of the middle and superior frontal gyri and precuneus (more prominent onthe right side) were observed. Second, the semantic memory condition disclosed extensive activations in the medial and orbital frontal cortexbilaterally, the left angular gyrus, and predominantly the left anterior part of the middle temporal gyri. The findings from this study arediscussed in light of the available neuropsychological data obtained in brain-damaged subjects and functional neuroimaging studies.© 2003 Elsevier Inc. All rights reserved.

Keywords: PET; Music; Semantic memory; Episodic memory; Frontal lobes; HERA model

Introduction

Many moments of our life are associated with a song ora particular melody. However, the neural substrates of mu-sical perception and musical memory processes are stilllittle known. Although it is common belief that perceptionof music is a specific ability of the right hemisphere, studiesin brain-damaged subjects demonstrate that musical capa-bilities are distributed in both hemispheres (Lechevalier etal., 1995; Peretz, 1994, 2001; Platel, 2002). Thus, in the

brain there is no “center for music” but a number of neuralnetworks that deal with the different components of musicperception (e.g., pitch, timber, rhythm, intensity); these neu-ral networks are very sensitive to musical training andexpertise and could differ from one subject to another (Al-tenmuller, 2001; Schlaug, 2001). Over the past 10 years, themost relevant information relating to the musical processeshave clearly emerged from functional neuroimaging studies,but most of these have addressed perceptual treatments(Zatorre et al., 1992; Sergent et al., 1992b; Platel et al.,1997; Griffiths et al., 1999). So far, very few neuropsycho-logical, psychophysical, or imaging studies have related tothe mnemonic aspects of music, and the concept of a “mod-ular” musical memory is little considered in the psycholog-ical and neuropsychological literatures.

* Corresponding author. EMI-E.0218 Inserm-Universite´ de Caen,U.F.R. de Psychologie, Universite´ de Caen, Esplanade de la Paix, 14032Caen Cedex, France.

E-mail address: [email protected] (H. Platel).

NeuroImage 20 (2003) 244–256 www.elsevier.com/locate/ynimg

1053-8119/03/$ – see front matter © 2003 Elsevier Inc. All rights reserved.doi:10.1016/S1053-8119(03)00287-8

Experimental psychology and psychoacoustic investiga-tions mainly relate to the elementary characteristics of mu-sical perception and more rarely on the mnemonic processesas such, while the rare studies that did so focussed onworking rather than long-term memory processes (Deutsch,1970; Semal and Demany, 1991, 1993). Experimental in-vestigations of the recall of well-known or new musicalparts generally concurred in showing the importance ofmelodic relative to rhythmic information in both the encod-ing and the retrieval processes for musical material (Halp-ern, 1984; Dowling et al., 1995; Hebert and Peretz, 1997).This supports the idea of a melodic advantage in the con-stitution of the musical lexicon.

Findings in brain-damaged subjects suggest that musicalmemory could have a different cerebral organization fromother kinds of memory, such as verbal and visual memory.Clinical reports show that although identification and rec-ognition of a musical piece appears to involve both hemi-spheres, integrity of the left hemisphere is however critical(see review in Lechevalier et al., 1995), as illustrated by aleft brain-damaged patient who exhibited impaired melodicidentification despite intact melodic discrimination (Eu-stache et al., 1990). Interestingly, these disturbances inmelodic identification were found to be dissociated from thelanguage impairment observed following cerebral lesionsinvolving specifically the verbal semantic processes (Signo-ret et al., 1987). In addition, the studies carried out byZatorre (1985) and Samson and Zatorre (1991) in patientswith right or left temporal lobectomies revealed a major roleof the right temporal areas in the recognition of unknownmelodies, suggesting that such recognition, which is notbased on a semantic strategy, is subtended by a perceptiveanalysis and a comparison of the melodies. The observationof patient C.N. (Peretz, 1996) is particularly relevant be-cause, several years after having sustained bilateral tempo-ral lesions, this patient was unable to memorize new musicaltunes. Furthermore, there was a lack of priming effect formusical material in this patient, which suggested abolishedaccess and encoding specific to music. For the author, thisclinical case argues for the existence of a long-term memorysubsystem specific to musical material.

Another argument in favour of a possible separate long-term memory subsystem between language and musiccomes from event-related brain potentials studies (ERPs;Besson and Schon, 2001), which show differential ERPeffects for semantic processes when subjects focus theirattention only to the lyrics or to the music of opera excerpts.

The aim of this work was to determine the neural substratesunderlying the semantic and episodic components of musicmemory. The distinction between episodic and semanticmemory has become very popular since it was first proposedby Tulving in 1972. Episodic memory is conceived as thememory of life events linked to their spatial and temporalcontext of acquisition, while semantic memory corresponds tothe memory of concepts, transcending a particular context(Tulving, 1985, 2001). In this work, we define musical seman-

tic memory as that referring to “well-known” excerpts of musicstored in memory without it being possible to retrieve thetemporal or spatial circumstances surrounding their encounter.Semantic memory allows us to identify or to have a strongfeeling of knowing for familiar songs or melodies. We mayname the tune (composer or performer) or just have the capac-ity to hum or whistle the subsequent notes of a melody. Mu-sical semantic memory may represent a musical lexicon, sep-arate of a verbal lexicon, even though strong links certainlyexist between them. Our previous PET study, which dealt withthe music components of music perception, did tackle thesemantic processing of music (Platel et al., 1997). In thisinvestigation, judging whether or not musical sequences werefamiliar induced specific activation of the left inferior frontalgyrus (area 47 of Brodmann) and anterior part of the leftsuperior temporal gyrus relative to attention to the other mu-sical components (i.e., rhythm, pitch, and timber). However, intheir study of mental imagery for familiar melodies, Halpernand Zatorre (1999) claimed that musical semantic memoryprimarily engaged right-sided regions. These findings howeverappear somewhat to conflict with the large neuroimaging lit-erature about recall of verbal or visual material from semanticmemory (see Cabeza and Nyberg, 2000, for review), in whichpreferential implication of the left temporal and prefrontalregions is a regular feature. It may therefore be argued that theright-sided engagement reported by Halpern and Zatorre(1999) may reflect the nonverbal nature of the musical materialused by these authors. The neural substrates of musical seman-tic memory therefore remain uncertain.

Episodic memory for musical information will be re-ferred here as the capacity to recognize a musical excerpt(whether familiar or not) for which the spatiotemporal con-text surrounding its former encounter (i.e., when, where,and how) can be recalled. Neuroimaging studies have con-firmed the importance of the medial temporal and frontallobes in the encoding and retrieval of episodic information(Cabeza and Nyberg, 2000; Fletcher and Henson, 2001).Mayes and Montaldi (2001) however note that functionalneuroimaging has been much less successful at confirmingthe roles of the midline diencephalon and basal forebrainstructures in episodic memory tasks. Explanatory hypothe-ses stressed both technical and methodological limitations(Desgranges et al., 1998). Yet, other structures, such as theanterior cingulate and the precuneus, have been found toengage during episodic memory tasks (Sanders et al., 2000).Concerning the prefrontal lobes, the HERA model (Tulvinget al., 1994) ascribes to the left prefrontal cortex a prefer-ential role in the encoding process of episodic material andthe recall of semantic information, while the right prefrontalcortex would preferentially operate in the recall of episodicinformation. Nyberg (1998) further proposed that the rightanterior prefrontal cortex would be involved in all memorytasks, and the right posterior prefrontal cortex in the moredifficult retrieval conditions. Other studies (Kelley et al.,1998) have suggested that lateralization of activations maydepend not so much on whether encoding or retrieval are

245H. Platel et al. / NeuroImage 20 (2003) 244–256

primarily being engaged, but on whether verbal or hard-to-verbalize materials are being processed. So far, however,experimental data have almost exclusively concerned verbalor visual material, and practically nothing is known aboutthe functional anatomy of episodic memory for music.

Based on our previous PET study (Platel et al., 1997), aswell as on the neuropsychological literature, we hypothesizethat musical semantic processes will induce activation of theleft inferior frontal and left anterior temporal areas, butcould also produce right prefrontal activations (Halpern andZatorre, 1999). Concerning musical episodic memory, weanticipate activation of the classic episodic memory net-work, namely involving the right prefrontal cortex, the an-terior cingulate gyrus, the precuneus, and potentially alsothe hippocampal regions, while whether or not additionalactivation specifically related to the use of musical materialwould occur cannot be firmly predicted.

Materials and methods

Subjects

Nine young healthy men were selected from a populationof university students. They were all right-handed and freeof any psychiatric or organic pathology and had normalhearing. To avoid specific cognitive strategies related tomusical expertise, these subjects were selected so as tobelong to class I of Wertheim and Botez (1959), i.e., “Mu-sical people without theoretical musical studies and musicalknowledge.” We purposely studied nonmusical subjects sothat our findings would generalize to the largest population,whereas musicians have specific strategies for listening tomusic, which depend in particular on the played instrument.The subjects who participated in this research were selectedon the basis of two principal criteria: first, they were to be“common listeners” (i.e., not music lovers, who tend tolisten to a specific type of music only), and second, theywere to have normal performances in a test of pitch percep-tion. The subjects selected for the PET study were verysimilar to the subjects who took part in the selection of thefamiliar and nonfamiliar melodies; thus, the general musicalculture of these two groups was equivalent.

They were told that the experiment in which they were totake part related to the perception of the music, but neverinformed that the experiment related to musical memory, sothat they did not train themselves to memorize melodiesbefore the experiment. All of them gave written informedconsent prior to participation and the research protocol wasapproved by the regional ethics committee. The study wasperformed in line with the Declaration of Helsinki.

Nature of the musical material

Musical material was specially created for this study andcomprised 128 short melodies (5 s) played without orches-

tration, but with the same timber of instrument (flute). Allwere real melodies, extracted from the classic and modernrepertoires, but excluding songs so as to limit verbal asso-ciations. We also excluded extracts which might spontane-ously evoke autobiographical memories, such as the “wed-ding march” or melodies used in popular TV spots. The 128tunes comprised 64 “ familiar” and 64 “nonfamiliar” tunes.The familiar melodies were those judged very familiar bymore than 70% of subjects in a pilot study of 150 subjectsmatched with the experimental sample,1 while the nonfa-miliar melodies were those judged as unknown by morethan 80% of the subjects from the same population. Bothsamples were extracted from a larger database of melodiesthat was used in this preexperimental study.

Paradigm

In the musical semantic memory task, the instruction wasto classify the melody heard as familiar or nonfamiliar. Halfof the stimuli were “ familiar” based on the above criteria(target items), and half were “nonfamiliar.” According toour design (see below), this semantic judgment will involveincidental encoding of the stimuli, and the subjects will beasked to recall the latter in the subsequent episodic memorytasks. We deliberately eschewed explicit encoding in ourparadigm so as to avoid the subjects from doing a doubletask, namely, memorizing the items while judging theirfamiliarity. To highlight activations specific to musical se-mantic memory, a perceptive control condition was de-signed where decisional and motor processes were present.The instruction was to indicate if the last two notes of eachsequence had the same pitch. Since by necessity the seman-tic task contained both familiar and nonfamiliar items, twodistinct control tasks had to be constructed, one containingonly familiar and the other only nonfamiliar melodies. Noneof the melodies used in these two perceptual control taskswere employed in the other conditions. So as to equalizeattentional involvement, these control tasks were made to bedifficult, with hit rates ranging from 60% to 70% in preex-perimental studies.

As already explained above, musical episodic memorywas studied by asking the subjects to recognize, amongdistractors, melodies (either familiar or nonfamiliar) that

1 Some examples of selected familiar melodies: L. W. Beethoven,excerpts from “Symphony Nos. 6 and 9” ; D. Brubeck, “Take Five” ; M.Mussorgsky, “Pictures at an exhibition” ; A. Vivaldi, excerpts from “TheSeasons” ; P.I. Tchaikovsky excerpts from “Nutcracker Suite” ; J. Williams,“Raiders March” ; S. Prokofiev, excerpts from “Peter and the Wolf” ; A.Dvorak, “Symphony No. 9, from the New World” ; M. Ravel, “Daphnis etChloe” ; W.A. Mozart, “Eine Kleine Nachtmusik” ; Vangelis, “Conquest ofParadise” ; E. Grieg, excerpts from “Peer Gynt Suite.” Some examples ofselected nonfamiliar melodies: L. W. Beethoven, excerpts from “Corolian”overture, “Piano sonata No. 14” ; W.A. Mozart, “Salzburg symphony No.1” ; R. Wagner “Tannhausser Overture” ; C. Debussy, “Syrinx” ; Vangelis,“Antartica” ; P.I. Tchaikovsky, excerpts from “Swan Lake” ; B. Smetana,“The Moldau” ; F. Schubert, excerpts from “Symphony No. 6 and 8.”

246 H. Platel et al. / NeuroImage 20 (2003) 244–256

they heard earlier, during the semantic task. Because, incontrast with the semantic task, there was no obligation forthe episodic task to mix familiar and unfamiliar items and tobe consistent with the control tasks, two episodic tasks wereconstructed (i.e., using nonfamiliar and familiar items, to bereferred to as F and NF in what follows). In each, half of theitems were targets (previously heard melodies), and halfwere distractors. Thus, both episodic tasks had to be givenafter the semantic task.

Brain activations specifically related to musical episodicretrieval were to be assessed by comparing the episodictasks (NF � F) to the control tasks (NF � F), as for thesemantic task. In addition, separating the familiar and non-familiar items was meant to enable us to assess the effect of“ familiarity,” by comparing each episodic task (NF or F) tothe corresponding control task (NF or F) and each episodiccondition to one another (episodic NF vs. F, and F vs. NF).To highlight possible automatic semantic processes duringthe episodic and control tasks with familiar melodies (F), wecontrasted these conditions with the nonfamiliar episodic orcontrol conditions (NF). Moreover, we directly comparedepisodic (NF � F) versus semantic and semantic versusepisodic (NF � F) conditions, and finally the pattern ofactivations for control tasks (NF � F) was revealed bycomparing these control conditions with rest measurements.

Altogether therefore, five tasks were constructed, i.e.,one semantic, two episodic, and two control tasks, with, asstated above, the semantic task always coming before theepisodic tasks. Because our preexperimental studies hadshown that the recall performance for the familiar items was

significantly better than that for nonfamiliar items, (�80and �70%, respectively), we partly counterbalanced thisdifference by giving the episodic task with nonfamiliaritems before that with familiar items, so as to lengthen theretention interval for the familiar items. In addition, theitems of the semantic tasks of each version were refreshedbetween the scans, before the episodic tasks (see Fig. 1).

In all five conditions, the subjects were instructed torespond by pressing with their right index finger the “yes”or “no” button of a computer mouse. Each task lasted 2 minand consisted of eight target and eight nontarget items. Eachitem was � 5 s long, with an interstimulus interval of 3 s.Each subject carried out each task twice and thus twoversions of each task, using different items, were con-structed. The order of the two versions of each task wascounterbalanced. Finally, two “ rest” scans (eyes closed andwithout auditory stimulation) were acquired. Altogether,therefore, 12 PET scans were acquired per subject (i.e., 10activation tasks and two rest measurements). With a total ofnine subjects, we thus collected 18 scans per condition. Theentire sequence of scans is illustrated in Fig. 1.

Data acquisition and analysis

Data acquisition

Measurements of regional distribution of radioactivitywere performed with an ECAT HR� (Siemens) PET cam-era with full-volume acquisition allowing the reconstruction

Fig. 1. Experimental paradigm.


of 63 planes. Transmission scans were obtained with a 68Gasource prior to emission scans. The duration of each scanwas 90 s. About 7 mCi of H2O15 was administered as a slowbolus in the left antecubital vein by means of an automaticinfusion pump. Each experimental condition was started30 s before data acquisition and continued until scan com-pletion. This process was repeated for each of the 12 scans,for a total injected dose of �80 mCi. The interval betweeninjections was 6 min 40 s. Subjects were scanned whilelying supine in a darkened and quiet room. The head wasgently immobilized in a dedicated head rest. Head positionwas aligned transaxially to the orbitomeatal line with a laserbeam. The position of the head was controlled with the laserbeam prior to each injection.

Data analysis

All calculations and image transformation were per-formed on Unix System work stations. First, the 12 scans ofeach subject were realigned to each other, using the AIR3.0software. For subsequent data analysis, the Statistical Para-metric Mapping software (SPM99, Wellcome Departmentof Cognitive Neurology) implemented in the MATLABenvironment was used. The images were nonlinearly trans-formed into standard space, MRI template of SMP99, whichis in Talairach space. The images were smoothed using a12-mm Gaussian filter. The images were scaled to an overallCBF grand mean of 50 ml/100 g/min; we therefore refer to“adjusted rCBF” in what follows. We used a gray matterthreshold of 80% of the whole brain mean; and covariateswere centered before inclusion in the design matrix. AnANCOVA (analysis of covariance), using global activity asa confounding covariate, was performed on a pixel-by-pixelbasis. The results of t statistic (SPM {t}) were then trans-formed into a normal standard distribution (SPM {z}). Thesignificance cutoff was set at P � 0.05, cluster-level-cor-rected for multiple comparisons.

Anatomical/cytoarchitechtonic localization of the signif-icant activations was based on the SPM99 MRI templateand Talairach’s coordinates, which were obtained using M.Brett’s linear transforms (see http://www.mrc-cbu.cam-.ac.uk/Imaging/mnispace.html). All the coordinates listed inthe sections below are SPM99 coordinates.

Results

Behavioral data

The average performance across the nine subjects wasconsistent with that obtained in the preexperimental popu-lation, with nearly 70% hit rate in the control tasks, morethan 80% in the semantic task, and 89% in the episodic taskwith familiar melodies (Fig. 2). As expected, the perfor-mance for the episodic task with nonfamiliar melodies (66%hit rate) was significantly lower than that for the familiar

melodies (P � 0.001). However, this result is particularlyrelated to the bad performance of one subject of the groupwho did not succeed in recognizing more than two nonfa-miliar items, whereas his performance for the familiar itemswas accurate. If this subject is discarded, the average per-centage of success becomes 73%, consistent with our stan-dards. Mean false alarms was rather small for all targetitems of the semantic and episodic tasks (�20%); as ex-pected, false alarms on nontargets items were commoner(mean 31%) for episodic tasks with nonfamiliar melodies.At debriefing, no subject expressed to have been aware ofthe dichotomy of the tasks according to the material (i.e.,familiar or not). More precisely, the subjects stated not tohave clearly realized that there were only familiar or un-known melodies for the control as for the episodic tasks.They did not try to make particular effort to maintain inmemory the material presented. There was no significantimprovement of the performance in the course of time.

PET data

Comparing the semantic with the control tasks (NF � F)showed extensive activation of the medial frontal areas (BA11 and 10) bilaterally, left middle temporal gyrus (extend-ing into the inferior frontal gyrus), and left angular gyrus(Table 1 and Fig. 3). Activation of these areas was notobserved in the right hemisphere.

Comparing the episodic (NF � F) with the control(NF � F) tasks revealed bilateral activation of the precu-neus (BA 7), the middle frontal gyri (BA 9 and 10), and themedial surface of the frontal lobe at the junction betweenBA9 and the anterior cingulate cortex. Although bilateral,these activations were stronger on the right hemisphere(Table 1 and Fig. 4).

Comparing the episodic (NF � F) with the semantictasks revealed activation of the precuneus (BA 7; right sidepredominantly), right superior frontal gyrus (BA 11), andright middle frontal gyrus (BA 8 and 9). All of these acti-vations were clearly right-sided (Table 1 and Fig. 5).

Comparing the semantic with the episodic tasks (NF �F) revealed activation of the bilateral medial frontal cortex(BA 11/10), left inferior and bilateral middle temporal gyrus(BA 20/21), and right cerebellum (Table 1 and Fig. 6).

Comparing the two control tasks (NF � F) with restrevealed extensive bilateral activation mainly of the lateraltemporal areas (middle and superior temporal gyri), bilat-eral inferior frontal gyrus, and cerebellum (Table 1 and Fig.7). The temporal activations included the primary and sec-ondary auditory cortices and were particularly conspicuouson the right hemisphere. Activation of the cerebellum, onthe other hand, was more pronounced on the left side.

Comparing the control tasks (F) with control tasks (NF)revealed activations of left middle and inferior frontal gyrus(BA 11/47), left precentral gyrus (BA 6, including Broca),and left medial frontal cortex (BA 8) (Table 1 and Fig. 8).

Comparing the episodic (F) with the control (F) tasks


http://www.mrc-cbu.cam.ac.uk/Imaging/mnispace.html

http://www.mrc-cbu.cam.ac.uk/Imaging/mnispace.html

revealed activation of the right precuneus (BA 7) and rightsuperior frontal gyrus (BA 10) only (Table 1 and Fig. 9).

Comparing the episodic (NF) with the control (NF) tasksrevealed bilateral activation of the superior and middlefrontal gyri (BA 10) bilateraly and of the medial frontalcortex (BA 8 and 9) (Table 1 and Fig. 10).

Finally, no significant activation was found for the control(NF) versus control (F) contrast and for the two direct com-parison between the episodic tasks (NF vs. F or F vs. NF).

Discussion

The activation pattern of the control perceptive tasks (con-trol NF � F vs. rest) is an awaited result and consolidates theidea of a right temporal preferential contribution in the percep-tion of melody and pitch judgment (Zatorre et al., 1994). Inaddition to temporal activations, bilateral activation of theintrasylvian (opercular) inferior frontal areas is observed. Asshown in Fig. 7 (axial cuts), these frontal activations do notreach the outer surface of the cortex and are not observed in thecomparisons relating to the memory tasks. These frontal acti-vations could correspond to working memory processes orcontribute to the perceptual analysis of the melodies (Fletcherand Henson, 2001). Activations of the cerebellum reflect par-

ticularly the motor control, and these were more accentuated inthe left hemisphere because the task response was carried outwith the right-hand fingers. The performance for these controltasks indicates that these tasks were difficult. However, theycould be performed, according to debriefing, without a feelingof failure, consistent with our goal. Accordingly, compared tothe rest condition, the pattern of activation mainly reflected anauditory perceptual activity. Consequently, we consider thatour control tasks were appropriate perceptual control condi-tions to serve as reference for memory tasks.

Compared to the control tasks, the activation patternsobserved for the semantic and episodic tasks were clearlyindependent (Figs. 3 and 4). Only a very limited overlap ofactivation may be considered in the medial frontal regions.These distinct and new results, obtained here with musicalmaterial, confirm the significant functional independencebetween semantic and episodic memory processes, alreadydocumented with verbal and visuospatial material (Cabezaand Nyberg, 2000; Mayes and Montaldi, 2001).

Semantic memory

During the semantic task, activation was found to beextensive in two distinct brain regions: first, the medialfrontal region (involving mainly BA 11 and 10) and, sec-

Fig. 2. Hit rates for the memory tasks and the control tasks: Semantic, semantic tasks carried out during PET scans; Episodic NF, episodic tasks withnon-familiar melodies; Episodic F, episodic tasks with familiar melodies; Control NF, perceptive/control tasks with nonfamiliar melodies; Control F,perceptive/control tasks with familiar melodies.


Table 1Activation results

t (z) Cluster size x, y, z

Semantic vs. control (NF � F)Bilateral medial frontal cortex (BA 11/10) 7.03 (6.29) 6197 0, 59, 08

6.14 (5.62) �4, 17, �166.00 (5.51) 6, 15, �18

Left middle temporal gyrus (BA 21) 6.14 (5.61) 2932 �55, �9, �155.43 (5.05) �57, �27, �2

Left inferior frontal gyrus (BA 47) 5.54 (5.14) �38, 13, �16Left angular gyrus (BA 39) 4.93 (4.63) 220 �51, �59, 29

Episodic (NF � F) vs. control (NF � F)Right precuneus (BA 7) 5.96 (5.47) 782 2, �62, 33Right middle frontal gyrus (BA 10) 5.67 (5.24) 1825 30, 58, �3

4.97 (4.67) 34, 62, 11Left middle frontal gyrus (BA 10) 5.23 (4.89) 964 �34, 56, �8

4.17 (3.99) �28, 62, 22Medial frontal cortex (BA 9) 3.99 (3.83) 439 2, 36, 22

Episodic (NF � F) vs. semanticRight precuneus (BA 7/BA 19) 4.57 (4.71) 637 36, �66, 38Precuneus (BA 7) 4.50 (4.27) 373 4, �56, 42Right superior frontal gyrus (BA 11) 4.80 (4.53) 558 34, 52, �14Right middle frontal gyrus (BA 8/9) 4.35 (4.14) 450 38, 12, 44

Semantic vs. episodic (NF � F)Bilateral medial frontal cortex (BA 11/10) 6.32 (5.75) 3252 0, 60, 10

5.39 (5.02) �4, 18, �18Right cerebellum 4.59 (4.35) 484 32, �84, �38Right middle temporal gyrus (BA 21) 4.54 (4.31) 363 56, 4, �24Left inferior and middle temporal gyri (BA 20/21) 3.84 (3.69) 544 �48, �26, �22

3.79 (3.65) �54, �2, �18Control (NF � F) vs. rest

Right superior temporal gyrus (BA 22) 12.37 (inf) 3922 63, �21, 59.6 (inf) 53, �10, �3

Left superior temporal gyrus (BA 22) 7.54 (6.65) 2788 �59, �17, 36.73 (6.06) �51, �2, �7

Left cerebellum 7.11 (6.34) 6688 �38, �67, �206.90 (6.19) �36, �60, �26

Right cerebellum 6.09 (5.58) 32, �73, �18Left thalamus 6.76 (6.09) 3314 �14, �17, 17Left caudate 5.75 (5.31) �30, 25, 02Left inferior frontal gyrus (BA 44) 5.48 (5.09) �44, 3, 20Right inferior frontal gyrus (BA 45) 4.91 (4.62) 772 40, 22, 10

4.36 (4.15) 34, 20, 17Right superior frontal gyrus (BA 11) 4.82 (4.54) 403 22, 56, �16

4.74 (4.48) 26, 49, �23Control (F) vs. control (NF)

Left middle and inferior frontal gyrus (BA 11/47) 5.52 (5.13) 875 �38, 50, �204.00 (3.83) �42, 24, �8

Left precentral gyrus (BA 6, Broca area) 4.84 (4.56) 1157 �42, 0, 364.33 (4.12) �54, 12, 6

Medial frontal cortex (BA 8) 4.37 (4.16) 403 �10, 36, 40Episodic (F) vs. control (F)

Right precuneus (BA 7) 5.45 (5.07) 551 5, �67, 33Right superior frontal gyrus (BA 10) 4.58 (4.34) 270 28, 57, 0

Episodic (NF) vs. control (NF)Right superior and middle frontal gyri (BA 10) 5.18 (4.85) 1242 32, 59, �16

4.37 (4.16) 38, 57, 6Left superior and middle frontal gyri (BA 10) 5.08 (4.77) 961 �32, 54, �8

4.26 (4.06) �24, 57, 10Medial frontal cortex (BA 8/9) 4.95 (4.65) 704 10, 33, 48

4.34 (4.13) �4, 33, 33

Note. Areas significantly activated at P � 0.001, but using the P � 0.05 corrected for multiple comparisons (cluster level) cutoff. Anatomical localizationof the significant activations and approximate Brodmann’s areas was as described under Materials and methods. Stereotaxic coordinates shown are those listedin SPM99.


Fig. 3. Semantic tasks versus control tasks (NF � F). Significantly activated regions at the threshold of P � 0.05 corrected for multiple comparisons,displayed with surface rendering. Illustrated here is the relative contribution of the different conditions of our paradigm, according to the “effects of interests”for selected peaks. The contrast was centred around zero, and the ordinate of each plot is the mean size of the effect for each condition � SEM, within thepeak voxel.Fig. 4. Episodic tasks (NF � F) versus control tasks (NF � F). Same as Fig. 3.


ond, the left hemisphere involving an extensive band of themiddle temporal gyrus (BA 21) and extending into theinferior frontal gyrus (BA 47). Further to these two mainactivation foci, an isolated activation of the left angulargyrus was noted. The semantic versus episodic contrast(Fig. 6), however revealed an additional right middle tem-poral gyrus activation (BA 21), suggesting a contribution ofthese right-sided areas in semantic memory processes. This

activation profile, mainly lateralized to the left hemisphere,is entirely consistent with previous functional neuroimagingof semantic memory tasks (Cabeza and Nyberg, 2000;Fletcher and Henson, 2001) and also with the HERA model(Tulving et al., 1994). The diagrams showing the effects ofinterests (Fig. 3) clearly document no noticeable participa-tion of the two control and episodic conditions in the findingof activation in these medial frontal and left middle tempo-

Fig. 5. Episodic tasks (NF � F) versus semantic tasks. Significantly activated regions at the threshold of P � 0.05 corrected for multiple comparisons,displayed with surface rendering.Fig. 6. Semantic tasks versus episodic tasks (NF � F). Significantly activated regions at the threshold of P � 0.05 corrected for multiple comparisons,displayed with surface rendering.Fig. 7. Control tasks (NF � F) versus rest. Significantly activated regions at the threshold of P � 0.05 corrected for multiple comparisons displayed withsurface rendering (left) and illustrative cuts (right) of the SPM99 T1 weights MRI template.Fig. 8. Control tasks (F) versus control tasks (NF). Significantly activated regions at the threshold of P � 0.05 corrected for multiple comparisons, displayedwith surface rendering.


ral regions. On the other hand, there was a substantial levelof activity of these regions during rest, suggesting a heter-ogeneous cognitive activity during this condition among thesubjects and justifying the importance of designing controlconditions that strongly engage attention, as purposely im-plemented for this paradigm.

Taken together, these data confirm a functional asymme-try in favour of the left hemisphere for semantic memorytasks. Regarding the medial frontal activation, similar find-ings have been reported occasionally with semantic tasks(Sergent et al., 1992a; Kapur et al., 1994; Vandenberghe etal., 1996). According to the review of Cabeza and Nyberg(2000), this activity would be specifically linked to theprocess of categorization of semantic information. Wetherefore consider that in our study, the medial frontalactivation would specifically reflect the categorization pro-cess (i.e., the decision on familiar vs. nonfamiliar melodies).For example, the results obtained by Tempini et al. (1998)are in close agreement with the present investigation. Theirstudy, in which the association of names and familiar faces(famous people) was studied, revealed an activation of themedial frontal areas (BA 10 and 11), left anterior temporalareas (BA 21), and angular gyrus. The activation of the leftangular gyrus may contribute to this categorization processand also possibly to attempts in verbal labeling of the heardmelodies. An activation of the left angular gyrus has alsobeen observed previously in tasks of verb or word genera-tion (Frith et al., 1991; Warburton et al., 1996).

Thus, activation of neither the medial frontal areas northe left angular gyrus appears to be very specific to musicalsemantic memory. In contrast, could the activation of theleft middle temporal and inferior frontal regions character-ize the access to a musical semantic memory? This latterfinding replicates that we obtained previously in a task ofjudgment of musical familiarity (Platel et al., 1997). Fur-

thermore, activation of the left inferior frontal gyrus, al-though less extensive than in our earlier study, was obtainedhere with a different musical material and experimentalparadigm. Additional elements support the hypothesis thatthis activation points to cortical regions that underlie musi-cal semantic memory. One meta-analysis of activationpeaks from PET studies relating to the perception of lan-guage and music (data from Petersen et al., 1988, andSergent et al., 1992b) revealed an absence of completeoverlap of activations (Drury and Van Essen, 1997). Thus,the perception of simple sounds, melodies, or timber pro-duces mainly temporal and prefrontal activations that onlypartially overlap those obtained for the perception of pho-nemes, logatoms, or real words (Petersen et al., 1988; Ser-gent et al., 1992b; Demonet et al., 1994; Zatorre et al., 1992;Platel et al., 1997), and activation peaks obtained withmusical material appeared more anteriorly located than forlinguistic material. We suggest therefore that the left ante-rior temporal and inferior frontal regions subtend specificnonverbal auditory processing abilities and that they mightunderlie a musical lexicon.

Why should there be a left hemisphere specialization formusical semantic memory? The prevalent idea that the righthemisphere preferentially underlies the perception of musicarises primarily from experimental studies (in particularwith the dichotic listening technique) using nonfamiliarmelodies (Peretz, 1994). Meanwhile, the few neuropsycho-logical case studies in which a specific impairment of mu-sical identification was present (in the absence of perceptualdeficit) almost exclusively concerned left hemisphere le-sions (Dupre and Nathan, 1911; Souques and Baruk, 1930;Eustache et al., 1990). The few functional neuroimagingstudies that have focused on musical memory (Zatorre et al.,1994; Matteis et al., 1997; Holcomb et al., 1998; Halpernand Zatorre, 1999) showed mainly bilateral, but preferen-

Fig. 9. Episodic tasks (F) versus control tasks (F). Comparison for only familiar melodies. Significantly activated regions at the threshold of P � 0.05corrected for multiple comparisons, displayed in SPM99 standard cuts (sagittal, coronal, and transverse).Fig. 10. Episodic tasks. (NF) versus control tasks (NF). Comparison for only nonfamiliar melodies. Significantly activated regions at the threshold of P �0.05 corrected for multiple comparisons, displayed in SPM99 standard cuts (sagittal, coronal, and transverse).


tially right-sided, frontal and anterior temporal activations.However, apart from the work of Halpern and Zatorre(1999), these studies did not address musical semanticmemory, but rather recognition processes for nonfamiliarmelodies. In the Halpern and Zatorre study (1999), thecomparison of the second task (to imagine the continuationof familiar melodies) with the third task (mental repetitionof new melodies) was designed to remove the functionalactivity related to auditory analysis, working memory, andmental imagery and to reveal only the recovery processes insemantic memory. This comparison revealed activation inthe right inferior and middle frontal gyri, the right inferiorand superior temporal gyri, the right anterior cingulate, andthe parietal cortex. Activation of the left inferior and middlefrontal gyri was also observed, though with less intensitythan on the right side. According to Halpern and Zatorre, theobserved asymmetry in favor of the right hemisphere wouldbe related to the musical specificity of the cognitive pro-cesses. There might however be some difficulty in the in-terpretation of this finding, given a contamination by epi-sodic processes, acknowledged by the authors in theirparadigm. Indeed, in contrast to our study, their subjectswere selected initially so as to have a strong feeling offamiliarity with the chosen musical extracts, and moreover,each subject was trained to imagine the continuation ofthese familiar melodies before the scan sessions. In contrast,our paradigm was specifically designed for the study ofmusical semantic memory, and we believe that the prefer-ential left-sided activation pattern obtained for the semantictasks does not only reflect verbal processing during perfor-mance of this task. Indeed, all the familiar melodies wereselected so as not to be popular songs (see Materials andmethods) and to be difficult to name for nonmusicians. Forexample, an extract like the “Toccata and Fugue in DMinor” from J.S. Bach would be easily judged as familiarby virtually all nonmusicians subjects, but only few of themwould be able to produce the title and/or the composer ofthis musical piece.

Thanks to dichotomizing the familiar and nonfamiliarmaterial in our design, it was also possible, in addition to thesemantic condition, to assess the brain activity involved insemantic memory during the control tasks with familiarmelodies. In other words, contrasting the control tasks withfamiliar versus nonfamiliar melodies makes it possible tohighlight the cerebral areas dedicated to the implicit accessto musical semantic memory. This contrast (Fig. 8) revealedsignificant differences in the left prefrontal areas (in partic-ular BA 6), as well as the left middle, inferior, and medialfrontal regions. Thus, a left dominance was again presentfor this contrast, strengthening the idea that mainly leftcerebral areas support the access to musical semantic mem-ory. However, activations observed in this comparisoncould also reflect processes of humming, if subjects invol-untarily sung (covertly) those melodies that were familiar tothem.

In agreement with Scott et al. (2000), we think that the

right posterior superior temporal regions play a specific partin the dynamic treatment of pitch variations (for music andlanguage) and that the left hemisphere homologous areasare more particularly dedicated to the phonological treat-ment and the comprehension of language. However, basedon the present work and our earlier study (Platel et al.,1997), we believe that the anterior part of the left temporalcortex is particularly involved in nonverbal semantic pro-cesses and could sustain musical semantic representations.Overall, therefore, the left middle temporal activations ob-served in our semantic task would mainly represent theaccess to musical semantic memory while the frontal acti-vations would represent essentially the process of categori-zation of the presented melodies.

Episodic memory

Recognition of the familiar and nonfamiliar melodiesheard at the time of the semantic task induced a profile ofactivation clearly different from that obtained for the se-mantic task. The observed activation of the rostral-most partof the middle frontal areas (BA 9 and 10) and the precuneus(BA 7), although bilateral, predominantly involved the righthemisphere (Fig. 4). Predominantly right-sided frontal andprefrontal activation has been frequently reported with re-trieval of either verbal or nonverbal material (Cabeza andNyberg, 2000). Comparing the episodic versus semantictasks (Fig. 5) confirmed first the right dominance for theretrieval processes and second that our musical semanticmemory task mobilized no evident resource in the righthemisphere given that subtracting the semantic processesfrom the episodic tasks did not decrease or remove theseactivations of the right hemisphere.

Regarding activation of the precuneus, although previousfunctional imaging studies on perception and processing ofmusical material have reported activation near this area(Sergent et al., 1992b; Platel et al., 1997), similar findingshave concerned nonmusical material, and thus this findinghas been interpreted as reflecting a process of mental im-agery triggered by the particular task (Kosslyn et al., 1997).However, precuneus activation has also been observed inepisodic memory tasks, regardless of whether or not theitems possessed imageable characteristics (Krause et al.,1999). According to Kapur et al. (1995), this cerebral regionwould be particularly involved in episodic retrieval andmore precisely in the success of episodic recall. This inter-pretation would fit well with our results, as our musicalmaterial did not involve particularly imageable features.Further arguments in favor of this interpretation would bethe lack of activation of the precuneus in our semantic taskdespite use of the same musical material as for the episodictask and the fact that, at debriefing, no subject mentionedhaving employed a specific mental representation strategyfor episodic recall, whether the melodies were familiar ornot. In addition, the separate comparisons between the ep-isodic and the control tasks for familiar (Fig. 9) and nonfa-


miliar melodies (Fig. 10) show that the recognition of thefamiliar melodies is associated with strictly right-lateralizedactivations (precuneus and superior frontal regions),whereas the recognition of nonfamiliar melodies producesbilateral activations of the frontal areas (medial, superior,and middle). Since our subjects could easily recognize thefamiliar melodies, these data would support the idea that theprecuneus activation was linked with the success of epi-sodic retrieval. On the other hand, the significant engage-ment of the frontal regions during the recognition of thenonfamiliar melodies reflects undoubtedly the difficultiesexperienced by the subjects to achieve this task and suggeststhe occurrence of executive control processes (such asmaintenance, planning, and inhibition). Nolde et al. (1998)also proposed that the extent of left prefrontal activationsduring retrieval probably increases as the executive de-mands of retrieval increase.

Overall, the results of the present investigation would bein agreement with the HERA model proposed by Tulvingand collaborators (1994), which is based on functionalasymmetry in favor of the left hemisphere for semanticmemory research and right hemisphere dominance for epi-sodic retrieval. Despite the fact that the HERA model re-mains controversial owing to conflicting results from somefunctional neuroimaging studies, the review of the literaturecarried out by Cabeza and Nyberg (2000) highlighted thefact that the majority of the published data reported suchmore or less marked asymmetry (Blanchet et al. 2001). Thebilaterality of frontal activations observed in many studiesusing widely different paradigms does however mitigate thenotion of a strict right-sided predominance for episodicretrieval. Some works suggest notably that lateralization ofactivations may not depend so much on whether encodingor retrieval are primarily being engaged, but more onwhether verbal or hard-to-verbalize materials are being pro-cessed (Kelley et al., 1998; Mayes and Montaldi, 2001).This hypothesis fits poorly with our results, showing moreleft-sided engagement of frontal regions for episodic re-trieval of hard-to-verbalize (nonfamiliar) melodies and clearright-sided activations with easier retrieval of familiar mel-odies. Our feeling, which concurs with that of Nolde et al.(1998), is that the extent of left prefrontal activations duringretrieval probably reflects the executive demands.

To conclude, these new findings from a study of seman-tic and episodic memory of musical material are altogetherconsistent with the results established earlier with verbal orvisuospatial material. However, some functional specificityfor musical memory does appear to exist. Based on our data,the left anterior temporal cortex would appear particularlyinvolved in semantic memory for musical material.

Acknowledgments

We thank the staff of the MRC Cyclotron Unit for scanningfacilities. This work was supported by Inserm U.320.

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