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S l t Mt A A ti ti i Di fl P ti S l t Mt A A ti ti i Di fl P ti Supplementary Motor Area Activation in Disfluency Perception Supplementary Motor Area Activation in Disfluency Perception Supplementary Motor Area Activation in Disfluency Perception. Supplementary Motor Area Activation in Disfluency Perception. A fMRI St d f Li t N lR t S t l A fMRI St d f Li t N lR t S t l An fMRI Study of Listener Neural Responses to Spontaneously An fMRI Study of Listener Neural Responses to Spontaneously An fMRI Study of Listener Neural Responses to Spontaneously An fMRI Study of Listener Neural Responses to Spontaneously P d d U fill d d Fill d P P d d U fill d d Fill d P Produced Unfilled and Filled Pauses Produced Unfilled and Filled Pauses Produced Unfilled and Filled Pauses Produced Unfilled and Filled Pauses Robert Eklund Robert Eklund 1 & Martin Ingvar Martin Ingvar 23 23 Robert Eklund Robert Eklund 1 & Martin Ingvar Martin Ingvar 2, 3 2, 3 1 Department of Culture and Communication Linköping University Linköping Sweden Department of Culture and Communication Linköping University Linköping Sweden Department of Culture and Communication, Linköping University, Linköping, Sweden Department of Culture and Communication, Linköping University, Linköping, Sweden 2 Department of Clinical Neuroscience Karolinska Institute Stockholm Sweden Department of Clinical Neuroscience Karolinska Institute Stockholm Sweden 2 Department of Clinical Neuroscience, Karolinska Institute, Stockholm, Sweden Department of Clinical Neuroscience, Karolinska Institute, Stockholm, Sweden 3 Osher Center for Integrative Medicine Karolinska Institute Stockholm Sweden Osher Center for Integrative Medicine Karolinska Institute Stockholm Sweden Osher Center for Integrative Medicine, Karolinska Institute, Stockholm, Sweden Osher Center for Integrative Medicine, Karolinska Institute, Stockholm, Sweden ABSTRACT ABSTRACT INTRODUCTION INTRODUCTION METHOD METHOD (1) ABSTRACT ABSTRACT INTRODUCTION INTRODUCTION METHOD METHOD (1) Spontaneously produced Unfilled Pauses (UPs) and Almost no one is completely fluent when speaking Subjects Spontaneously produced Unfilled Pauses (UPs) and Filled Pauses (FPs) were played to subjects in an fMRI Almost no one is completely fluent when speaking. The subjects were 16 healthy adults (9 F/7M) ages 2254 (mean age 40.3, Filled Pauses (FPs) were played to subjects in an fMRI experiment. Speech disfluency has been studied since at least the standard deviation 9.5) with no reported hearing problems. All subjects h h dd d d b h db h dd experiment. Speech disfluency has been studied since at least the 1930s. Disfluencies have commonly been regarded as were right-handed as determined by the Edinburgh Handedness Inventory (Oldfield 1971) All subjects possessed higher education Following a For both stimuli increased activity was observed in the 1930s. Disfluencies have commonly been regarded as “performance errors” in speech production. (Oldfield, 1971). All subjects possessed higher education. Following a description of the study, including a description of fMRI methodology, For both stimuli increased activity was observed in the Primary Auditory Cortex (PAC) However FPs but not performance errors in speech production. written and informed consent was obtained from all subjects. A small ti i ti ti l d iit d Primary Auditory Cortex (PAC). However, FPs, but not UPs elicited modulation in the Supplementary Motor However, several studies indicate that certain kinds participation remuneration was also administered. UPs, elicited modulation in the Supplementary Motor Area (SMA) Brodmann Area 6 of disfluencies can have beneficial effects on listener Equipment Area (SMA), Brodmann Area 6. perception (Fraundorf & Watson, 2011; Barr & The fMRI scanner used was the General Electric 1.5T Excite HD Twinspeed t K li k I tit t MR t St kh l S d Th il Our results provide neurocognitive confirmation of the Seyfeddinipur, 2010; Ferreira & Bailey, 2004; Fox Tree, scanner at Karolinska Institute, MR -center, Stockholm, Sweden. The coil used was a General Electrics Standard bird-cage head-coil (1 5T) Our results provide neurocognitive confirmation of the alleged difference between FPs and other kinds of 2001, 1995; Reich, 1980). used was a General Electrics Standard bird cage head coil (1.5T). Sti l data alleged difference between FPs and other kinds of speech disfluency and could also provide a partial Stimulus data h l d d f h h h dl speech disfluency and could also provide a partial explanation for the previously reported beneficial The most common voiced disfluency is the filled pause The stimulus data used were excerpts from the humanhuman dialog speech data described in Eklund (2004: 187190) Subjects were asked to explanation for the previously reported beneficial effect of FPs on reaction times in speech perception (FP), “eh” . The reported average frequency of filled speech data described in Eklund (2004: 187190). Subjects were asked to play the role of travel agents listening to customers making travel bookings effect of FPs on reaction times in speech perception. pauses (FPs) ranges from 1.9 to 7 .6% at word level over the telephone, following a task sheet (Eklund, 2004: 185) Our results also have potential implications for two (Eklund, 2010). From the original data set, four speakers were chosen (2M/2F) and a Our results also have potential implications for two of the suggested functions of FPs: the “floor holding” number of sentences were excised that were fluent except that they included a number of UPs and an approximately equal number of FPs The of the suggested functions of FPs: the floor-holding and the “help me out” hypotheses Neurocognitive studies on disfluency have traditionally included a number of UPs and an approximately equal number of FPs. The resulting number of both UPs and FPs roughly corresponded to reported and the “help-me-out” hypotheses. consisted of electrophysiological studies, starting with resulting number of both UPs and FPs roughly corresponded to reported incidence of UPs and FPs in spontaneous speech. Keywords: speech disfluency filled pauses unfilled Kutas and Hillyard (1980) on the N400 component. Stimulus data are shown in Table 1 below. Keywords: speech disfluency, filled pauses, unfilled pauses speech perception spontaneous speech fMRI pauses, speech perception, spontaneous speech, fMRI, Auditory Cortex PAC Supplementary Motor Area SMA The present study uses functional Magnetic Resonance Auditory Cortex, PAC, Supplementary Motor Area, SMA, Brodmann Area 6 BA6 Imaging (fMRI) to analyse the effect of authentic Brodmann Area 6, BA6 disfluencies proper to study the effect of unfilled and filler pauses on brain processing. 1 2 3 1 2 3 METHOD METHOD (3) METHOD METHOD (2) COMMENTS COMMENTS ON ON DATA DATA AND AND ANALYSES ANALYSES 1 W d fMRI t td di fl ti t Post-experiment interview Experimental design 1. We used fMRI to study disfluency perception, not EEG ith it f t l t f h After the scanning, all subjects were interviewed in order to confirm that the had been a ake and foc sed d ing the e pe iment A self ating The stimulus files described above were used in an event-related i t Aft i iti l l li t i l i i th f EEG with its focus on temporal aspects of speech ti they had been awake and focused during the experiment. A self -rating scale of how attentive the subjects felt they had been during the sessions experiment. After initial localizer anatomical scanning sessions, the four stimulus files (M/F/M/F) were played During the intermissions the subjects perception. scale of how attentive the subjects felt they had been during the sessions was also used. All subjects reported that they had been attentive at a stimulus files (M/F/M/F) were played. During the intermissions the subjects were briefed whether they were still awake/focused on the task. 2. We investigated perceptual modulation caused by satisfactory level. Interstimulus intervals were of sufficient duration so as to allow for reliable BOLD i iti FP d UP d ll d t i SPM d FPs proper, not their effect on ensuing items MRI scans BOLD acquisition. FPs and UPs were modelled as events in SPM and were convolved using the Haemodynamic Response Function (HRF) in SPM (words, phrases) or general cognitive processing. For each subject, a T1-weighted coronal spoiled gradient recalled (SPGR) convolved using the Haemodynamic Response Function (HRF) in SPM. Experimental setting 3 Unlike previous studies where the auditory stimuli image volume was obtained to serve as anatomical reference ( / / fl l l 3 ) Experimental setting The subjects lay supine/head first in the scanner with earplugs to protect 3. Unlike previous studies where the auditory stimuli often have been scripted laboratory speech we (TR/TE= 24.0/ 6.0 ms; flip angle 35°; voxel size = 1 × 1 × 1 mm 3 ). The subjects lay supine/head first in the scanner with earplugs to protect them from scanner noise and headphones with the sound data played to often have been scripted laboratory speech, we used ecologically valid stimulus data Moreover, BOLD-sensitized fMRI was carried out by using echo-planar imaging EPI using 32 axial slices (TR/TE 2500/40 ms flip 90 deg voxel them from scanner noise and headphones with the sound data played to them. The perceived sound level was quite sufficient and no subjects used ecologically valid stimulus data. imaging EPI using 32 axial slices (TR/TE=2500/40 ms, flip = 90 deg, voxel size = 3. 75 × 3.75 × 4 mm 3 ). reported having any problems hearing what was said. Head movement was constrained using foam wedges and/or tape ANALYSES ANALYSES AND AND RESULTS RESULTS (1) size 3. 75 × 3.75 × 4 mm ). In total T2*-weighted images were collected four sessions: (3m30s/80 constrained using foam wedges and/or tape. ANALYSES ANALYSES AND AND RESULTS RESULTS (1) In total, T2 -weighted images were collected four sessions: (3m30s/80 volumes; 2m22s/53 volumes; 1m33s/33 volumes; 3m05s/70 volumes). Experimental instructions Using Fluent Speech (FS) as the baseline condition, P t i The subjects were instructed to listen carefully to what was said, as if they were the addressed travel agent but that they were not expected to react the following three contrasts were analyzed: Post-processing Th i t d d l d i M tL b R2007 d were the addressed travel agent, but that they were not expected to react verbally to the utterances or say anything, only that they needed to pay (1) Filled Pauses > Fluent Speech The images were post-processed and analyzed using MatLab R2007a and SPM5 (Friston et al 2007) The images were realigned co-registered and verbally to the utterances or say anything, only that they needed to pay attention to the information provided by the clients. All subjects (1) Filled Pauses > Fluent Speech SPM5 (Friston et al. , 2007). The images were realigned, co registered and normalized to the EPI template image in SPM5 and finally smoothed using understood the instructions without any confusion. (2) Unfilled Pauses > Fluent Speech a FWHM (Full-Width Half Maximum) of 6 mm. Experimental design: data acquisition (3) Filled Pauses > Unfilled Pauses Regressors pertaining to subject head movement (3 translational and 3 (3) Filled Pauses > Unfilled Pauses rotational) were included as parameters of no-interest in the general linear model at the first level of analysis The results were calculated with a False Discovery model at the first level of analysis. N bj t ldd d t h d ti f th i Rate at p < 0.05 (Genovese, Lazar & Nichols, 2002) No subjects were excluded due to head motion or for any other image acquisition related causes Analyses were also carried out using the SPM with a cluster level threshold of 10 contiguous voxels. acquisition related causes. Analyses were also carried out using the SPM Anatomy Toolbox (Amunts, Schleicher & Zilles, 2007; Eickhoff et al., 2007, No activation in Brodmann Area 22 associated with 2006, 2005). No activation in Brodmann Area 22, associated with semantic processing was observed semantic processing, was observed. 4 5 6 DISCUSSION DISCUSSION AND AND CONCLUSIONS CONCLUSIONS ANALYSES ANALYSES AND AND RESULTS RESULTS (2) POTENTIAL POTENTIAL IMPLICATIONS IMPLICATIONS DISCUSSION DISCUSSION AND AND CONCLUSIONS CONCLUSIONS ANALYSES ANALYSES AND AND RESULTS RESULTS (2) POTENTIAL POTENTIAL IMPLICATIONS IMPLICATIONS (1) UPs and FPs modulated the Primary Auditory Results are shown in Table 2 and Figures 13 below: (1) “floor-holding” hypothesis of FPs, as first (1) UPs and FPs modulated the Primary Auditory Cortex (PAC) That heightened attention Results are shown in Table 2 and Figures 13 below: T bl 2 L i f i ifi i i f h FDR d 0 05 d ih l (1) floor holding hypothesis of FPs, as first proposed by Maclay and Osgood (1959): that FPs Cortex (PAC). That heightened attention influences auditory perception has been shown Table 2. Locations of significant activation for three contrasts, FDR-corrected at p < 0.05 and with a cluster level threshold of 10 contiguous voxels, analyzed with SPM (Friston et al., 2007) and the SPM Anatomy proposed by Maclay and Osgood (1959): that FPs are used to keep the floor in polylogs influences auditory perception has been shown (Petkov et al 2004; and more) and the observed Toolbox (Eickhoff et al., 2007, 2006, 2007). Brodmann Areas were identified using the Talairach Atlas (Talairach & Tournoux, 1988) and the Talairach Client/Daemon (www .talairach.org), using a Cube Range are used to keep the floor in polylogs. O lt i l th t FP ld ibl b (Petkov et al. 2004; and more) and the observed attention-heightening function of FPs could setting of ±5 mm. Our results imply that FPs could possibly be t d ti f thi attention heightening function of FPs could possibly help explain the shorter reaction times to counter-productive for this. possibly help explain the shorter reaction times to linguistic stimuli that follow FPs as reported by e g (2) “help-me-out” hypothesis as suggested in linguistic stimuli that follow FPs as reported by e.g. Fox Tree (2001 1995) Note however that UPs (2) help-me-out hypothesis, as suggested in Clark and Wilkes Gibbs (1986): FPs are/can be Fox Tree (2001, 1995). Note, however, that UPs also exhibited PAC activation Clark and Wilkes-Gibbs (1986): FPs are/can be used as a (semi deliberate) signal asking for also exhibited PAC activation. used as a (semi-deliberate) signal asking for interlocutor help in conversation Both UPs and FPs lead to heightened attention. interlocutor help in conversation Our results imply that FPs might serve this (2) Filled Pauses activated (pre-)motor areas, function well. (e.g. BA6). (Not observed in UPs. ) K l d i Bik (1940) th t SMA i SUMMING SUMMING IT IT UP UP Known already since Brickner (1940) that SMA is i i h i f h d l SUMMING SUMMING IT IT UP UP Our results confessedly speculative suggest that active in the processing of speech, and several l d h f d b h d Our results confessedly speculative suggest that FPs but not FS/UPs activate motor areas in the later studies have confirmed both SMA and FPs, but not FS/UPs, activate motor areas in the listener brain pre-SMA play a role in both speech production listener brain. (e.g. Goldberg, 1985; Alario et al. , 2006) and Both FPs/UPs activate PAC which lends support to the speech perception (e.g. Iacoboni, 2008; Wilson et Both FPs/UPs activate PAC, which lends support to the attention heightening hypothesis that has been al. , 2004 and many more). attention-heightening hypothesis that has been forwarded in the literature and it would also seem FPs seem to kick -startspeech production forwarded in the literature and it would also seem clear that it is not the break in the speech stream per FPs seem to kick start speech production. clear that it is not the break in the speech stream per se that causes this activation since UPs seemingly do se that causes this activation, since UPs seemingly do not have this effect 7 8 9 not have this effect. 7 8 9

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Page 1: Sl tMtA AtitiiDifl P tiSupplementary Motor Area Activation ...robek28/pdf/Eklund_Ingvar_2016_Disfluency_… · Title: Microsoft PowerPoint - Interspeech2016_EklundIngvar_fMRI_POSTER_FINAL.ppt

S l t M t A A ti ti i Di fl P tiS l t M t A A ti ti i Di fl P tiSupplementary Motor Area Activation in Disfluency PerceptionSupplementary Motor Area Activation in Disfluency PerceptionSupplementary Motor Area Activation in Disfluency Perception.Supplementary Motor Area Activation in Disfluency Perception.pp y y ppp y y pA fMRI St d f Li t N l R t S t lA fMRI St d f Li t N l R t S t lAn fMRI Study of Listener Neural Responses to SpontaneouslyAn fMRI Study of Listener Neural Responses to SpontaneouslyAn fMRI Study of Listener Neural Responses to Spontaneously An fMRI Study of Listener Neural Responses to Spontaneously y p p yy p p y

P d d U fill d d Fill d PP d d U fill d d Fill d PProduced Unfilled and Filled PausesProduced Unfilled and Filled PausesProduced Unfilled and Filled PausesProduced Unfilled and Filled PausesRobert EklundRobert Eklund11 && Martin IngvarMartin Ingvar2 32 3Robert EklundRobert Eklund11 && Martin IngvarMartin Ingvar2, 32, 3gg

11 Department of Culture and Communication Linköping University Linköping SwedenDepartment of Culture and Communication Linköping University Linköping SwedenDepartment of Culture and Communication, Linköping University, Linköping, SwedenDepartment of Culture and Communication, Linköping University, Linköping, Sweden22 Department of Clinical Neuroscience Karolinska Institute Stockholm SwedenDepartment of Clinical Neuroscience Karolinska Institute Stockholm Sweden22 Department of Clinical Neuroscience, Karolinska Institute, Stockholm, SwedenDepartment of Clinical Neuroscience, Karolinska Institute, Stockholm, Sweden

33 Osher Center for Integrative Medicine Karolinska Institute Stockholm SwedenOsher Center for Integrative Medicine Karolinska Institute Stockholm SwedenOsher Center for Integrative Medicine, Karolinska Institute, Stockholm, SwedenOsher Center for Integrative Medicine, Karolinska Institute, Stockholm, Sweden

ABSTRACTABSTRACT INTRODUCTIONINTRODUCTION METHODMETHOD ((11))ABSTRACTABSTRACT INTRODUCTIONINTRODUCTION METHODMETHOD ((11))

Spontaneously produced Unfilled Pauses (UPs) and Almost no one is completely fluent when speaking SubjectsSpontaneously produced Unfilled Pauses (UPs) andFilled Pauses (FPs) were played to subjects in an fMRI

Almost no one is completely fluent when speaking.The subjects were 16 healthy adults (9 F/7M) ages 22–54 (mean age 40.3,Filled Pauses (FPs) were played to subjects in an fMRI

experiment. Speech disfluency has been studied since at least thej y ( / ) g ( g ,

standard deviation 9.5) with no reported hearing problems. All subjectsh h d d d d b h d b h d dexperiment. Speech disfluency has been studied since at least the

1930s. Disfluencies have commonly been regarded aswere right-handed as determined by the Edinburgh Handedness Inventory(Oldfield 1971) All subjects possessed higher education Following a

For both stimuli increased activity was observed in the1930s. Disfluencies have commonly been regarded as“performance errors” in speech production.

(Oldfield, 1971). All subjects possessed higher education. Following adescription of the study, including a description of fMRI methodology,For both stimuli increased activity was observed in the

Primary Auditory Cortex (PAC) However FPs but notperformance errors in speech production. p y, g p gy,

written and informed consent was obtained from all subjects. A smallti i ti ti l d i i t dPrimary Auditory Cortex (PAC). However, FPs, but not

UPs elicited modulation in the Supplementary Motor However, several studies indicate that certain kinds participation remuneration was also administered.UPs, elicited modulation in the Supplementary MotorArea (SMA) Brodmann Area 6

,of disfluencies can have beneficial effects on listener Equipment

Area (SMA), Brodmann Area 6.perception (Fraundorf & Watson, 2011; Barr & The fMRI scanner used was the General Electric 1.5T Excite HD Twinspeed

t K li k I tit t MR t St kh l S d Th ilOur results provide neurocognitive confirmation of the

p p ( , ;Seyfeddinipur, 2010; Ferreira & Bailey, 2004; Fox Tree,

scanner at Karolinska Institute, MR-center, Stockholm, Sweden. The coilused was a General Electrics Standard bird-cage head-coil (1 5T)Our results provide neurocognitive confirmation of the

alleged difference between FPs and other kinds ofy p , ; y, ; ,

2001, 1995; Reich, 1980).used was a General Electrics Standard bird cage head coil (1.5T).

Sti l dataalleged difference between FPs and other kinds ofspeech disfluency and could also provide a partial

00 , 995; e c , 980)Stimulus data

h l d d f h h h d lspeech disfluency and could also provide a partialexplanation for the previously reported beneficial The most common voiced disfluency is the filled pause The stimulus data used were excerpts from the human–human dialog

speech data described in Eklund (2004: 187–190) Subjects were asked toexplanation for the previously reported beneficialeffect of FPs on reaction times in speech perception

y p(FP), “eh”. The reported average frequency of filled

speech data described in Eklund (2004: 187–190). Subjects were asked toplay the role of travel agents listening to customers making travel bookingseffect of FPs on reaction times in speech perception. ( ), p g q y

pauses (FPs) ranges from 1.9 to 7.6% at word levelp y g g g gover the telephone, following a task sheet (Eklund, 2004: 185)

Our results also have potential implications for two

p ( ) g(Eklund, 2010). From the original data set, four speakers were chosen (2M/2F) and a

Our results also have potential implications for twoof the suggested functions of FPs: the “floor holding”

( , )number of sentences were excised that were fluent except that theyincluded a number of UPs and an approximately equal number of FPs Theof the suggested functions of FPs: the floor-holding

and the “help me out” hypothesesNeurocognitive studies on disfluency have traditionally included a number of UPs and an approximately equal number of FPs. The

resulting number of both UPs and FPs roughly corresponded to reportedand the “help-me-out” hypotheses. consisted of electrophysiological studies, starting withresulting number of both UPs and FPs roughly corresponded to reportedincidence of UPs and FPs in spontaneous speech.

Keywords: speech disfluency filled pauses unfilledKutas and Hillyard (1980) on the N400 component. Stimulus data are shown in Table 1 below.

Keywords: speech disfluency, filled pauses, unfilledpauses speech perception spontaneous speech fMRIpauses, speech perception, spontaneous speech, fMRI,Auditory Cortex PAC Supplementary Motor Area SMA

The present study uses functional Magnetic ResonanceAuditory Cortex, PAC, Supplementary Motor Area, SMA,Brodmann Area 6 BA6

Imaging (fMRI) to analyse the effect of authenticBrodmann Area 6, BA6

disfluencies proper to study the effect of unfilled andfiller pauses on brain processing.

11 22 3311 22 33

METHODMETHOD ((33))METHODMETHOD ((22)) COMMENTSCOMMENTS ONON DATADATA ANDAND ANALYSESANALYSESOO ((33))OO (( ))

1 W d fMRI t t d di fl ti tPost-experiment interviewExperimental design 1. We used fMRI to study disfluency perception, notEEG ith it f t l t f hAfter the scanning, all subjects were interviewed in order to confirm that

the had been a ake and foc sed d ing the e pe iment A self atingThe stimulus files described above were used in an event-related

i t Aft i iti l l li t i l i i th fEEG with its focus on temporal aspects of speech

tithey had been awake and focused during the experiment. A self-ratingscale of how attentive the subjects felt they had been during the sessions

experiment. After initial localizer anatomical scanning sessions, the fourstimulus files (M/F/M/F) were played During the intermissions the subjects

perception.scale of how attentive the subjects felt they had been during the sessionswas also used. All subjects reported that they had been attentive at a

stimulus files (M/F/M/F) were played. During the intermissions the subjectswere briefed whether they were still awake/focused on the task. 2. We investigated perceptual modulation caused by

satisfactory level.Interstimulus intervals were of sufficient duration so as to allow for reliableBOLD i iti FP d UP d ll d t i SPM d

g p p yFPs proper, not their effect on ensuing items

MRI scansBOLD acquisition. FPs and UPs were modelled as events in SPM and wereconvolved using the Haemodynamic Response Function (HRF) in SPM

p p , g(words, phrases) or general cognitive processing.

For each subject, a T1-weighted coronal spoiled gradient recalled (SPGR)convolved using the Haemodynamic Response Function (HRF) in SPM.

Experimental setting

( , p ) g g p g

3 Unlike previous studies where the auditory stimulij , g p g ( )image volume was obtained to serve as anatomical reference( / / fl l l 3)

Experimental setting

The subjects lay supine/head first in the scanner with earplugs to protect

3. Unlike previous studies where the auditory stimulioften have been scripted laboratory speech we(TR/TE= 24.0/ 6.0 ms; flip angle 35°; voxel size = 1 × 1 × 1 mm3).The subjects lay supine/head first in the scanner with earplugs to protect

them from scanner noise and headphones with the sound data played tooften have been scripted laboratory speech, weused ecologically valid stimulus dataMoreover, BOLD-sensitized fMRI was carried out by using echo-planar

imaging EPI using 32 axial slices (TR/TE 2500/40 ms flip 90 deg voxel

them from scanner noise and headphones with the sound data played tothem. The perceived sound level was quite sufficient and no subjects

used ecologically valid stimulus data.imaging EPI using 32 axial slices (TR/TE=2500/40 ms, flip = 90 deg, voxelsize = 3.75 × 3.75 × 4 mm3).

reported having any problems hearing what was said. Head movement wasconstrained using foam wedges and/or tape ANALYSESANALYSES ANDAND RESULTSRESULTS ((11))size 3.75 × 3.75 × 4 mm ).

In total T2*-weighted images were collected four sessions: (3m30s/80constrained using foam wedges and/or tape. ANALYSESANALYSES ANDAND RESULTSRESULTS ((11))

In total, T2 -weighted images were collected four sessions: (3m30s/80volumes; 2m22s/53 volumes; 1m33s/33 volumes; 3m05s/70 volumes).

Experimental instructions Using Fluent Speech (FS) as the baseline condition,; / ; / ; / )

P t iThe subjects were instructed to listen carefully to what was said, as if theywere the addressed travel agent but that they were not expected to react

the following three contrasts were analyzed:Post-processing

Th i t d d l d i M tL b R2007 d

were the addressed travel agent, but that they were not expected to reactverbally to the utterances or say anything, only that they needed to pay (1) Filled Pauses > Fluent SpeechThe images were post-processed and analyzed using MatLab R2007a and

SPM5 (Friston et al 2007) The images were realigned co-registered and

verbally to the utterances or say anything, only that they needed to payattention to the information provided by the clients. All subjects

(1) Filled Pauses > Fluent SpeechSPM5 (Friston et al., 2007). The images were realigned, co registered andnormalized to the EPI template image in SPM5 and finally smoothed using

understood the instructions without any confusion. (2) Unfilled Pauses > Fluent Speechp g y ga FWHM (Full-Width Half Maximum) of 6 mm.Experimental design: data acquisition

(3) Filled Pauses > Unfilled PausesRegressors pertaining to subject head movement (3 translational and 3

(3) Filled Pauses > Unfilled Pausesrotational) were included as parameters of no-interest in the general linearmodel at the first level of analysis

The results were calculated with a False Discoverymodel at the first level of analysis.

N bj t l d d d t h d ti f th iRate at p < 0.05 (Genovese, Lazar & Nichols, 2002)

No subjects were excluded due to head motion or for any other imageacquisition related causes Analyses were also carried out using the SPM with a cluster level threshold of 10 contiguous voxels.acquisition related causes. Analyses were also carried out using the SPMAnatomy Toolbox (Amunts, Schleicher & Zilles, 2007; Eickhoff et al., 2007, No activation in Brodmann Area 22 associated with2006, 2005). No activation in Brodmann Area 22, associated with

semantic processing was observedsemantic processing, was observed.

44 55 66

DISCUSSIONDISCUSSION ANDAND CONCLUSIONSCONCLUSIONSANALYSESANALYSES ANDAND RESULTSRESULTS ((22)) POTENTIALPOTENTIAL IMPLICATIONSIMPLICATIONSDISCUSSIONDISCUSSION ANDAND CONCLUSIONSCONCLUSIONSANALYSESANALYSES ANDAND RESULTSRESULTS ((22)) POTENTIALPOTENTIAL IMPLICATIONSIMPLICATIONS

(1) UPs and FPs modulated the Primary AuditoryResults are shown in Table 2 and Figures 1–3 below: (1) “floor-holding” hypothesis of FPs, as first(1) UPs and FPs modulated the Primary AuditoryCortex (PAC) That heightened attention

Results are shown in Table 2 and Figures 1–3 below:T bl 2 L i f i ifi i i f h FDR d 0 05 d i h l

(1) floor holding hypothesis of FPs, as firstproposed by Maclay and Osgood (1959): that FPsCortex (PAC). That heightened attention

influences auditory perception has been shownTable 2. Locations of significant activation for three contrasts, FDR-corrected at p < 0.05 and with a clusterlevel threshold of 10 contiguous voxels, analyzed with SPM (Friston et al., 2007) and the SPM Anatomy

proposed by Maclay and Osgood (1959): that FPsare used to keep the floor in polylogsinfluences auditory perception has been shown

(Petkov et al 2004; and more) and the observedToolbox (Eickhoff et al., 2007, 2006, 2007). Brodmann Areas were identified using the Talairach Atlas(Talairach & Tournoux, 1988) and the Talairach Client/Daemon (www.talairach.org), using a Cube Range

are used to keep the floor in polylogs.

O lt i l th t FP ld ibl b(Petkov et al. 2004; and more) and the observedattention-heightening function of FPs could

( , ) ( g), g gsetting of ±5 mm. Our results imply that FPs could possibly be

t d ti f thiattention heightening function of FPs couldpossibly help explain the shorter reaction times to

counter-productive for this.possibly help explain the shorter reaction times tolinguistic stimuli that follow FPs as reported by e g (2) “help-me-out” hypothesis as suggested inlinguistic stimuli that follow FPs as reported by e.g.Fox Tree (2001 1995) Note however that UPs

(2) help-me-out hypothesis, as suggested inClark and Wilkes Gibbs (1986): FPs are/can beFox Tree (2001, 1995). Note, however, that UPs

also exhibited PAC activationClark and Wilkes-Gibbs (1986): FPs are/can beused as a (semi deliberate) signal asking foralso exhibited PAC activation. used as a (semi-deliberate) signal asking forinterlocutor help in conversation Both UPs and FPs lead to heightened attention. interlocutor help in conversation

Our results imply that FPs might serve this(2) Filled Pauses activated (pre-)motor areas, function well.

(e.g. BA6). (Not observed in UPs.)( g ) ( )

K l d i B i k (1940) th t SMA iSUMMINGSUMMING ITIT UPUP

Known already since Brickner (1940) that SMA isi i h i f h d l

SUMMINGSUMMING ITIT UPUPOur results confessedly speculative suggest thatactive in the processing of speech, and several

l d h f d b h dOur results – confessedly speculative – suggest thatFPs but not FS/UPs activate motor areas in thelater studies have confirmed both SMA and FPs, but not FS/UPs, activate motor areas in thelistener brainpre-SMA play a role in both speech production listener brain.

(e.g. Goldberg, 1985; Alario et al., 2006) andBoth FPs/UPs activate PAC which lends support to thespeech perception (e.g. Iacoboni, 2008; Wilson et Both FPs/UPs activate PAC, which lends support to theattention heightening hypothesis that has beenal., 2004 and many more). attention-heightening hypothesis that has beenforwarded in the literature and it would also seem

FPs seem to “kick-start” speech productionforwarded in the literature and it would also seemclear that it is not the break in the speech stream per FPs seem to kick start speech production. clear that it is not the break in the speech stream perse that causes this activation since UPs seemingly dose that causes this activation, since UPs seemingly donot have this effect

77 88 99not have this effect.

77 88 99