Tʼainʼt What You Say, Itʼs the Way That You Say It—Left Insula - IA de Investigación especializada en el MIT

Tʼainʼt What You Say, Itʼs the Way That You Say It—Left Insula
and Inferior Frontal Cortex Work in Interaction with
Superior Temporal Regions to Control the
Performance of Vocal Impersonations

Carolyn McGettigan1,2, Frank Eisner3, Zarinah K. Agnew1, Tom Manly4,
Duncan Wisbey1, and Sophie K. Scott1

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Abstracto

■ Historically, the study of human identity perception has
focused on faces, but the voice is also central to our expressions
and experiences of identity [Belin, PAG., Fecteau, S., & Bedard, C.
Thinking the voice: Neural correlates of voice perception. Trends
in Cognitive Sciences, 8, 129–135, 2004]. Our voices are highly
flexible and dynamic; talkers speak differently, depending on
their health, emotional state, and the social setting, así como
extrinsic factors such as background noise. Sin embargo, hasta la fecha,
there have been no studies of the neural correlates of identity
modulation in speech production. In the current fMRI experi-
mento, we measured the neural activity supporting controlled
voice change in adult participants performing spoken impres-

siones. We reveal that deliberate modulation of vocal identity
recruits the left anterior insula and inferior frontal gyrus, apoyo-
ing the planning of novel articulations. Bilateral sites in posterior
superior temporal/inferior parietal cortex and a region in right
middle/anterior STS showed greater responses during the emula-
tion of specific vocal identities than for impressions of generic
accents. Using functional connectivity analyses, we describe roles
for these three sites in their interactions with the brain regions
supporting speech planning and production. Our findings mark
a significant step toward understanding the neural control of
vocal identity, with wider implications for the cognitive control
of voluntary motor acts. ■

INTRODUCCIÓN

Voices, like faces, express many aspects of our identity
(Belin, Fecteau, & Bedard, 2004). From hearing only a
few words of an utterance, we can estimate the speakerʼs
gender and age, their country or even specific town of
birth, as well as more subtle evaluations on current mood
or state of health (Karpf, 2007). Some of the indexical
cues to speaker identity are clearly expressed in the
voice. The pitch (or fundamental frequency, F0) del
voice of an adult male speaker tends to be lower than
that of adult women or children, because of the thicken-
ing and lengthening of the vocal folds during puberty in
human men. The secondary descent of the larynx in adult
men also increases the spectral range in the voice, reflejar-
ing an increase in vocal tract length.

Sin embargo, the human voice is also highly flexible, y
we continually modulate the way we speak. The Lombard
efecto (Lombard, 1911) describes the way that talkers
automatically raise the volume of their voice when the

1Institute of Cognitive Neuroscience, University College London,
2Department of Psychology, Royal Holloway, Universidad de
Londres, 3Max Planck Institute for Psycholinguistics, Nimega,
Los países bajos, 4MRC Cognition and Brain Sciences Unit,
Cambridge, Reino Unido

© 2013 Massachusetts Institute of Technology Published under a
Creative Commons Attribution-NonCommercial 3.0 no portado
(CC BY-NC 3.0) licencia.

auditory environment is perceived as noisy. In the social
context of conversations, interlocutors start to align their
behaviors, from body movements and breathing patterns
to pronunciations and selection of syntactic structures
(Pardo, 2006; Garrod & Pickering, 2004; McFarland,
2001; Chartrand & Bargh, 1999; Condon & Ogston,
1967). Laboratory tests of speech shadowing, where par-
ticipants repeat speech immediately as they hear it, tener
shown evidence for unconscious imitation of linguistic and
paralinguistic properties of speech (Kappes, Baumgaertner,
Peschke, & Ziegler, 2009; Shockley, Sabadini, & Fowler,
2004; Bailly, 2003). Giles and colleagues (Giles, Coupland,
& Coupland, 1991; Giles, 1973) put forward the Com-
munication Accommodation Theory to account for pro-
cesses of convergence and divergence in spoken language
pronunciation—namely, they suggest that talkers change
their speaking style to modulate the social distance be-
tween them and their interlocutors, with convergence
promoting greater closeness. It has been argued by others
that covert speech imitation is central to facilitating com-
prehension in conversation (Pickering & Garrod, 2007).
Aside from these short-term modulations in speech,
changes in vocal behavior can also be observed over much
longer periods—the speech of Queen Elizabeth II has
shown a gradual progression toward standard southern

Revista de neurociencia cognitiva 25:11, páginas. 1875–1886
doi:10.1162/jocn_a_00427

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British pronunciation (Harrington, Palethorpe, & watson,
2000).

Although modulations of the voice often occur outside
conscious awareness, they can also be deliberate. A re-
cent study showed that student participants could change
their speech to sound more masculine or feminine, por
making controlled alterations that simulated target-
appropriate changes in vocal tract length and voice pitch
(Cartei, Cowles, & Reby, 2012). En efecto, speakers can
readily disguise their vocal identity (sullivan & Schlichting,
1998), which makes forensic voice identification noto-
riously difficult (Eriksson et al., 2010; Ladefoged, 2003).
Notablemente, when control of vocal identity is compromised,
Por ejemplo, in Foreign Accent Syndrome (p.ej., Scott,
Clegg, Rudge, & Burgess, 2006), the change in the patientʼs
vocal expression of identity can be frustrating and debili-
tating. Interrogating the neural systems supporting vocal
modulation is an important step in understanding human
vocal expression, yet this dynamic aspect of the voice is a
missing element in existing models of speech production
(Hickok, 2012; Tourville & Guenther, 2011).

Speaking aloud is an example of a very well practised
voluntary motor act (Jurgens, 2002). Voluntary actions
need to be controlled in a flexible manner to adjust to
changes in environment and the goals of the actor. El
main purpose of speech is to perform the transfer of a
linguistic/conceptual message. Sin embargo, we control our
voices to achieve intended goals on a variety of levels,
from acoustic–phonetic accommodation to the auditory
ambiente (cocinero & Lu, 2010; Lu & cocinero, 2009) a
socially motivated vocal behaviors reflecting how we wish
to be perceived by others (Pardo, Gibbons, Suppes, &
Krauss, 2012; Pardo & Jay, 2010). Investigations of the
cortical control of vocalization have identified two neuro-
logical systems supporting the voluntary initiation of innate
and learned vocal behaviors, where expressions such as
emotional vocalizations are controlled by a medial frontal
system involving the ACC and SMA, whereas speech and
song are under the control of lateral motor cortices
( Jurgens, 2002). De este modo, patients with speech production
deficits following strokes to lateral inferior motor structures
still exhibit spontaneous vocal behaviors such as laughter,
crying, and swearing, despite their severe deficits in volun-
tary speech production (Groswasser, Korn, Groswasser-
Reider, & Solzi, 1988). Electrical stimulation studies
show that vocalizations can be elicited by direct stim-
ulation of the anterior cingulate (p.ej., laughter; descrito
by Sem-Jacobsen & Torkildsen, 1960) and lesion evidence
shows that bilateral damage to anterior cingulate prevents
the expression of emotional inflection in speech ( Jurgens
& por cramon, 1982).

In healthy participants, a detailed investigation of the
lateral motor areas involved in voluntary speech produc-
tion directly compared voluntary inhalation/exhalation
with syllable repetition. The study found that the func-
tional networks associated with laryngeal motor cortex
were strongly left-lateralized for syllable repetition but

bilaterally organized for controlled breathing (Simonyan,
Ostuni, Ludlow, & Horwitz, 2009). Sin embargo, that design
did not permit further exploration of the modulation of
voluntary control within either speech or breathing. Este
aspect has been addressed in a study of speech prosody,
which reported activations in left inferior frontal gyrus
(IFG) and dorsal premotor cortex for the voluntary mod-
ulation of both linguistic and emotional prosody, eso
overlapped with regions sensitive to the perception of
these modulations (Aziz-Zadeh, Sheng, & Gheytanchi,
2010).

Some studies have addressed the neural correlates of
overt and unintended imitation of heard speech (Reiterer
et al., 2011; Peschke, Ziegler, Kappes, & Baumgaertner,
2009). Peschke and colleagues found evidence for un-
conscious imitation of speech duration and F0 in a shad-
owing task in fMRI, in which activation in right inferior
parietal cortex correlated with stronger imitation of dura-
tion across participants. Reiterer and colleagues (2011)
found that participants with poor ability to imitate non-
native speech showed greater activation (and lower gray
matter density) in left premotor, inferior frontal, and in-
ferior parietal cortical regions during a speech imitation
tarea, compared with participants who were highly rated
mimics. The authors interpret this as a possible index of
greater effort in the phonological loop for less skilled imi-
tatores. Sin embargo, en general, the reported functional imag-
ing investigations of voluntary speech control systems
have typically involved comparisons of speech outputs
with varying linguistic content, Por ejemplo, conectado
speech of different linguistic complexities (Dhanjal,
Handunnetthi, patel, & Inteligente, 2008; Blank, Scott, Murphy,
Warburton, & Inteligente, 2002) or pseudowords of varying
length and phonetic complexity (Papoutsi et al., 2009;
Bohland & Guenther, 2006).

To address the ubiquitous behavior of voluntary mod-
ulation of vocal expression in speech, while holding the
linguistic content of the utterance constant, we carried
out an fMRI experiment in which we studied the neural
correlates of controlled voice change in adult speakers of
English performing spoken impressions. The participants,
who were not professional voice artists or impressionists,
repeatedly recited the opening lines of a familiar nursery
rhyme under three different speaking conditions: normal
voice (norte), impersonating individuals (I), and impersonat-
ing regional and foreign accents of English (A). The nature
of the task is similar to the kinds of vocal imitation used in
everyday conversation, Por ejemplo, in reporting the
speech of others during storytelling. We aimed to uncover
the neural systems supporting changes in the way speech
is articulated, in the presence of unvarying linguistic con-
tent. We predicted that left-dominant orofacial motor
control centers, including the left IFG, insula, and motor
corteza, as well as auditory processing sites in superior
temporal cortex, would be important in effecting change to
speaking style and monitoring the auditory consequences.
Beyond this, we aimed to measure whether the goal of

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the vocal modulation—to imitate a generic speaking style/
accent versus a specific vocal identity—would modulate
the activation of the speech production network and/or
its connectivity with brain regions processing information
relevant to individual identities.

MÉTODOS

Participantes

Twenty-three adult speakers of English (seven women;
edad media = 33 años 11 meses) were recruited who
were willing to attempt spoken impersonations. All had
healthy hearing and no history of neurological incidents
nor any problems with speech or language (self-reported).
Although some had formal training in acting and music,
none had worked professionally as an impressionist or voice
artist. The study was approved by the University College
London Department of Psychology Ethics Committee.

Design and Procedure

Participants were asked to compile in advance lists of 40 indi-
viduals and 40 accents they could feasibly attempt to imper-
sonate. These could include any voice/accent with which
they were personally familiar, from celebrities to family
miembros (p.ej., “Sean Connery,” “Carlyʼs Mum”). Asimismo,
the selected accents could be general or specific (p.ej.,
“French” vs. “Blackburn”).

Functional imaging data were acquired on a Siemens
Avanto 1.5-T scanner (Siemens AG, Erlangen, Alemania)
in a single run of 163 echo-planar whole-brain volumes
(repetition time = 8 segundo, acquisition time = 3 segundo, echo
time = 50 mseg, flip angle = 90°, 35 axial slices, 3 mm ×
3 mm × 3 mm in-plane resolution). A sparse-sampling
rutina (Edmister, Talavage, Ledden, & Weisskoff, 1999;
Hall et al., 1999) was employed, with the task performed
during a 5-sec silence between volumes.

Había 40 trials of each condition: normal voice
(norte), impersonating individuals (I), impressions of re-
gional and foreign accents of English (A), and a rest base-
line (B). The mean list lengths across participants were
36.1 (DE = 5.6) for condition I and 35.0 (DE = 6.9) para
A (a nonsignificant difference; t(1, 22) = .795, pag = .435).
When submitted lists were shorter than 40, some names/
accents were repeated to fill the 40 ensayos. Condition order
was pseudorandomized, with each condition occurring
once in every four trials. Participants wore electrodynamic
headphones fitted with an optical microphone (MR Confon
GmbH, Magdeburg, Alemania). Using MATLAB (Matemáticas,
Cª, Natick, MAMÁ) with the Psychophysics Toolbox exten-
sión (Brainard, 1997) and a video projector (Eiki Inter-
national, Cª, Rancho Santa Margarita, California), visual prompts
(“Normal Voice,” “Break” or the name of a voice/accent,
as well as a “Start speaking” instruction) were delivered
onto a front screen, viewed via a mirror on the head coil.
Each trial began with a condition prompt triggered by

the onset of a whole-brain acquisition. En 0.2 sec after
the start of the silent period, the participant was prompted
to start speaking and to cease when the prompt dis-
appeared (3.8 sec later). In each speech production trial,
participants recited the opening line from a familiar nurs-
ery rhyme, such as “Jack and Jill went up the hill,” and were
reminded that they should not include person-specific
catchphrases or catchwords. This controlled for the
linguistic content of the speech across the conditions.
Spoken responses were recorded using Audacity (audacity.
sourceforge.net). After the functional run, a high-resolution
T1-weighted anatomical image was acquired (HIRes
MP-RAGE, 160 sagittal slices, voxel size = 1 mm3). El
total time in the scanner was around 35 mín..

Acoustic Analysis of Spoken Impressions

Because of technical problems, auditory recordings were
only available for 13 Participantes. El 40 tokens from the
three speech conditions—Normal Voice, Impersonations,
and Accents—was entered into a repeated-measures
ANOVA with Condition as a within-subject factor for each
of the following acoustic parameters: (i) duración (segundo), (ii)
intensidad (dB), (iii) mean F0 (Hz), (iv) minimum F0 (Hz),
(v) maximum F0 (Hz), standard deviation of F0 (Hz), (vi)
spectral center of gravity (Hz), y (vii) spectral standard
desviación (Hz). Three Bonferroni-corrected post hoc
paired t tests compared the individual conditions. Mesa 1
illustrates the results of these analyses, y figura 1 illus-
trates the acoustic properties of example trials from each
speech condition (taken from the same participant).

fMRI Analysis

Data were preprocessed and analyzed using SPM5 (Well-
come Trust Centre for Neuroimaging, Londres, Reino Unido). Func-
tional images were realigned and unwarped, coregistered
with the anatomical image, normalized using parameters
obtained from unified segmentation of the anatomical
imagen, and smoothed using a Gaussian kernel of 8 mm
FWHM. At the first level, the condition onsets were mod-
eled as instantaneous events coincident with the prompt to
speak, using a canonical hemodynamic response function.
Contrast images were calculated to describe each of the
four conditions (norte, I, A and B), each speech condition com-
pared with rest (N > B, I > B, A > B), each impression
condition compared with normal speech (I > N, A > N),
and the comparison of impression conditions (I > A).
These images were entered into second-level, one-sample
t tests for the group analyses.

The results of the conjunction analyses are reported at
a voxel height threshold of p < .05 (corrected for family- wise error). All other results are reported at an uncor- rected voxel height threshold of p < .001, with a cluster extent correction of 20 voxels applied for a whole-brain α of p < .001 using a Monte Carlo simulation (with 10,000 McGettigan et al. 1877 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f / . t u s e r o n 1 7 M a y 2 0 2 1 ANOVA t Test N vs. V t Test N vs. A t Test V vs. A Table 1. Acoustic Correlates of Voice Change during Spoken Impressions Acoustic Parameter Mean Normal Mean Voices Mean Accents Duration (sec) 2.75 3.10 2.98 Intensity (dB) Mean F0 (Hz) Min F0 (Hz) Max F0 (Hz) SD F0 (Hz) Spec CoG (Hz) Spec SD (Hz) 47.4 155.9 94.4 625.0 117.3 2100 1647 51.3 207.2 104.9 667.6 129.9 2140 1579 51.3 186.3 102.1 628.5 114.7 2061 1553 F 9.96 49.25 24.11 3.71 1.28 1.62 0.38 2.24 Sig. .006 .000 .000 .039 .295 .227 .617 .128 t 3.25 10.15 5.19 2.20 1.31 1.26 Sig. .021 .000 .001 .144 .646 .694 t 3.18 7.62 4.87 2.18 0.10 Sig. .024 .000 .001 .149 1.00 .240 1.00 t Sig. 2.51 .081 0.88 1.00 3.89 .006 0.77 1.00 2.15 3.30 1.49 .158 .019 .485 0.37 1.00 1.17 .789 0.39 2.05 1.00 .188 0.89 1.00 F0 = fundamental frequency, SD = standard deviation, Spec = spectral, CoG = center of gravity. Significance levels are Bonferroni-corrected (see Methods), with significant effects shown in bold. iterations) implemented in MATLAB (Slotnick, Moo, Segal, & Hart, 2003). Conjunction analyses of second-level contrast images were performed using the null conjunction approach (Nichols, Brett, Andersson, Wager, & Poline, 2005). Using the MarsBaR toolbox (Brett, Anton, Valabregue, & Poline, 2002), spherical ROIs (4 mm radius) were built around the peak voxels—parameter estimates were extracted from these ROIs to construct plots of activation. A psychophysiological interaction (PPI) analysis was used to investigate changes in connectivity between the conditions I and A. In each participant, the time course of activation was extracted from spherical volumes of inter- est (4 mm radius) built around the superior temporal peaks in the group contrast I > A (right middle/anterior
STS: [54 −3 −15], right posterior STS: [57 −36 12], izquierda
posterior STS: [−45 −60 15]). A PPI regressor described
the interaction between each volume of interest and a
psychological regressor for the contrast of interest (I >
A)—this modeled a change in the correlation between
activity in these STS seed regions and the rest of the
brain across the two conditions. The PPIs from each seed
region were evaluated in a first-level model that included
the individual physiological and psychological time
courses as covariates of no interest. A random-effects,
one-sample t test assessed the significance of each PPI
en el grupo (voxelwise threshold: pag < .001, corrected cluster threshold: p < .001). Post hoc pairwise t tests using SPSS (version 18.0; IBM, Armonk, NY) compared condition-specific parameter esti- mates (N vs. B and I vs. A) within the peak voxels in the voice change conjunction ((I > norte) ∩ (A > N)). To maintain
independence and avoid statistical “double-dipping,” an
iterative, hold-one-out approach was used in which the
peak voxels for each participant were defined from a group
statistical map of the conjunction ((I > N) ∩ (A > N))
using the other 22 Participantes. These subject-specific
peak locations were used to extract condition-specific

parameter estimates from 4-mm spherical ROIs built
around the peak voxel (using MarsBaR). Paired t tests
were run using a corrected α level of .025 (to correct
for two tests in each ROI).

The anatomical locations of peak and subpeak voxels
(al menos 8 mm apart) were labeled using the SPM Anatomy
Toolbox (versión 18; Eickhoff et al., 2005).

RESULTS AND DISCUSSION

Brain Regions Supporting Voice Change

Areas of activation common to the three speech output
conditions compared with a rest baseline (B; (N > B) ∩
(I > B) ∩ (A > B)) comprised a speech production net-
work of bilateral motor and somatosensory cortex, SMA,

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Cifra 1. Examples of the phrase “Jack and Jill went up the hill”
spoken by a single participant in the conditions Normal Voice, Accents,
and Impersonations. Top: A spectrogram of frequency against time
(where darker shading indicates greater intensity). Middle: El
fundamental frequency (F0) profile across each utterance. Bottom:
The intensity contour in decibels.

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Cifra 2. Activations common
to the three speech conditions
(Normal, Impersonations,
and Accents) comparado con
a rest baseline (voxel height
threshold p < .05, FWE- corrected). Numbers indicate the z coordinate in Montreal Neurological Institute (MNI) stereotactic space. D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j / . t f u s e r o n 1 7 M a y 2 0 2 1 superior temporal gyrus (STG), and cerebellum (Figure 2 and Table 2; Simmonds, Wise, Dhanjal, & Leech, 2011; Tourville & Guenther, 2011; Tourville, Reilly, & Guenther, 2008; Bohland & Guenther, 2006; Riecker et al., 2005; Blank et al., 2002; Wise, Greene, Buchel, & Scott, 1999). Activation common to the voice change conditions (I and A) compared with normal speech ((I >

norte) ∩ (A > N)) was found in left anterior insula, extending
laterally onto the IFG (orbital and opercular parts) and on
the right STG (Cifra 3 and Table 3). Planned post hoc
comparisons showed that responses in the left frontal sites
were equivalent for impersonations and accents (two-
cola, paired t test; t(22) = −0.068, corrected p = 1.00) y
during normal speech and rest (t(22) = 0.278, corregido

Mesa 2. Activation Common to the Three Speech Output Conditions

Contrast

No. of Voxels

Region

All Speech > Rest ((N > B) ∩

(I > B) ∩ (A > B))

963

Left postcentral gyrus/STG/precentral gyrus

−48 −15

39 14.15 7.07

Coordinate

852

Right STG/precentral gyrus/postcentral gyrus

Left cerebellum (lobule VI)

Left SMA

63 −15

3 13.60 6.96

−24 −60 −18

7.88 5.38

−3 −3

7.77 5.34

Right cerebellum (lobule VI), right fusiform gyrus

12 −60 −15

7.44 5.21

Right/left calcarine gyrus

Left calcarine gyrus

Right lingual gyrus

Right area V4

Left calcarine gyrus

Left thalamus

Right calcarine gyrus

3 −93

7.41 5.19

−15 −93 −3

6.98 5.02

15 −84 −3

6.73 4.91

30 −69 −12

6.58 4.84

−9 −81

6.17 4.65

−12 −24 −3

6.15 4.64

15 −69

6.13 4.63

Conjunction null analysis of all speech conditions (Normal, Impersonations, and Accents) compared with rest. Voxel height threshold p < .05 (FWE- corrected). Coordinates indicate the position of the peak voxel from each significant cluster, in MNI stereotactic space. McGettigan et al. 1879 p = 1.00). The right STG, in contrast, was significantly more active during impersonations than accents (two- tailed, paired t test; t(22) = 2.69, Bonferroni-corrected p = .027) and during normal speech compared with rest (t(22) = 6.64, corrected p < .0001). Thus, we demon- strate a partial dissociation of the inferior frontal/insular and sensory cortices, where both respond more during impressions than in normal speech, but where the STG shows an additional sensitivity to the nature of the voice change task—that is, whether the voice target is asso- ciated with a unique identity. Acoustic analyses of the impressions from a subset of participants (n = 13) indicated that the conditions in- volving voice change resulted in acoustic speech signals that were significantly longer, more intense, and higher in fundamental frequency (roughly equivalent to pitch) than normal speech. This may relate to the right-lateralized temporal response during voice change, as previous work has shown that the right STG is engaged during judg- ments of sound intensity (Belin et al., 1998). The right temporal lobe has also been associated with processing nonlinguistic information in the voice, such as speaker identity (von Kriegstein, Kleinschmidt, Sterzer, & Giraud, 2005; Kriegstein & Giraud, 2004; von Kriegstein, Eger, Kleinschmidt, & Giraud, 2003; Belin, Zatorre, & Ahad, 2002; Belin, Zatorre, Lafaille, Ahad, & Pike, 2000) and emotion (Schirmer & Kotz, 2006; Meyer, Zysset, von Cramon, & Alter, 2005; Wildgruber et al., 2005), although these re- sults tend to implicate higher-order regions such as the STS. The neuropsychology literature has described the im- portance of the left IFG and anterior insula in voluntary speech production (Kurth, Zilles, Fox, Laird, & Eickhoff, 2010; Dronkers, 1996; Broca, 1861). Studies of speech production have identified that the left posterior IFG and insula are sensitive to increasing articulatory complex- ity of spoken syllables (Riecker, Brendel, Ziegler, Erb, & Ackermann, 2008; Bohland & Guenther, 2006), but not to the frequency with which those syllables occur in every- day language (Riecker et al., 2008), suggesting involvement in the phonetic aspects of speech output rather than higher- order linguistic representations. Ackermann and Riecker (2010) suggest that insula cortex may actually be associated with more generalized control of breathing, which could be voluntarily modulated to maintain the sustained and finely controlled hyperventilation required to produce con- nected speech. In finding that the left IFG and insula can influence the way we speak, as well as what we say, we have also shown that they are not just coding abstract linguistic Figure 3. Brain regions supporting voice change. Bar plots show parameter estimates extracted from spherical ROIs centered on peak voxels. Annotations show the results of planned paired-sample t tests (two-tailed, with Bonferroni correction; *p < .05, ***p < .0001, ns = nonsignificant). Coordinates are in MNI space. D o w n l o a d e d f r o m l l / / / / j f / t t i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j . f / t u s e r o n 1 7 M a y 2 0 2 1 1880 Journal of Cognitive Neuroscience Volume 25, Number 11 Table 3. Neural Regions Recruited during Voice Change (Null Conjunction of Impersonations > Normal and Accents > Normal)

Contrast

No. de
Voxels

Region

Impressions > Normal Speech

180

LIFG (pars orb., pars operc.)/insula

((I > N) ∩ (A > N))

Left temporal pole

Right thalamus

Right STG

Right hippocampus

Left thalamus

Left hippocampus

Right insula

Right STG

Left hippocampus

Right STG

Right temporal pole

Left hippocampus

Right caudate nucleus

Left cerebellum (lobule VI)

Coordinate

−33

−54

−12

−27

−15

−24

−15

−24

−6

−24

−45

−6

−21

−3

−39

−9

−42

−60

−3

−9

−15

−6

−18

8.39

5.56

7.48

7.44

7.30

7.17

7.11

6.80

6.65

6.59

6.45

6.44

6.42

6.30

6.20

6.10

5.22

5.21

5.15

5.10

5.07

4.94

4.87

4.85

4.78

4.77

4.71

4.66

4.62

Voxel height threshold p < .05 (FWE-error corrected). Coordinates indicate the position of the peak voxel from each significant cluster, in MNI stereotactic space. LIFG = left IFG; pars orb. = pars orbitalis; pars operc. = pars opercularis. elements of the speech act. In agreement with Ackermann and Riecker (2010), we suggest that these regions may also play a role in more general aspects of voluntary vocal con- trol during speech, such as breathing and modulation of pitch. In line with this, our acoustic analysis shows that both accents and impressions were produced with longer durations, higher pitches, and greater intensity, all of which are strongly dependent on the way that breathing is controlled (MacLarnon & Hewitt, 1999, 2004). Effects of Target Specificity: Impersonations versus Accents A direct comparison of the two voice change conditions (I > A) showed increased activation for specific imper-
sonations in right middle/anterior STS, bilateral posterior
STS extending to angular gyrus (AG) on the left, and pos-
terior midline sites on cingulate cortex and precuneus
(Cifra 4 and Table 4; the contrast A > I gave no signifi-
cant activations). Whole-brain analyses of functional con-
nectivity revealed areas that correlated more positively
with the three sites on STS during impersonations than
during accents (Cifra 5 and Table 5). Strikingly, all three
temporal seed regions showed significant interactions
with areas typically active during speech perception and

tu
s
mi
r

oh
norte

1
7

METRO
a
y

2
0
2
1

Cifra 4. Greater activation for the production of specific
impersonations (I) than for accents (A). Coordinates are in MNI space.
Voxel height threshold p < .001, cluster threshold p < .001 (corrected). McGettigan et al. 1881 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t . f / Table 4. Brain Regions Showing Greater Activation during Specific Impersonations Contrast No. of Voxels Region Impersonation > Accents

Right STS

Left STS

Left middle cingulate cortex

Right STG

Coordinate

−45

−6

−3

−60

−48

−36

−15

5.79

4.62

4.48

4.35

4.46

3.82

3.73

3.66

Voxel height threshold p < .001 (uncorrected), cluster threshold p < .001 (corrected). Coordinates indicate the position of the peak voxel from each significant cluster, in MNI stereotactic space. D o w n l o a d e d f r o m production, with notable sites of overlap in sensorimotor lobules V and VI of the cerebellum and left STG. How- ever, there were also indications of differentiation of the three connectivity profiles. The left posterior STS seed region interacted with a speech production network including bilateral pre/postcentral gyrus, bilateral STG, and cerebellum (Price, 2010; Bohland & Guenther, 2006; Blank et al., 2002), as well as left-lateralized areas of anterior insula and posterior medial planum tempor- ale. In contrast, the right anterior STS seed interacted with the left opercular part of the IFG and left SMA, and the right posterior STS showed a positive interaction with the left inferior frontal gyrus/sulcus, extending to the left frontal pole. Figure 5 illustrates the more anterior dis- tribution of activations from the right-lateralized seed regions and the region of overlap from all seed regions in cerebellar targets. Our results suggest that different emphases can be distinguished between the roles performed by these superior temporal and inferior parietal areas in spoken impressions. In a meta-analysis of the semantic system, Binder, Desai, Graves, and Conant (2009) identified the AG as a high-order processing site performing the re- trieval and integration of concepts (Binder et al., 2009). The posterior left STS/AG activation has been implicated in the production of complex narrative speech and writing (Brownsett & Wise, 2010; Awad, Warren, Scott, Turkheimer, & Wise, 2007; Spitsyna, Warren, Scott, Turkheimer, & Wise, 2006) and, along with the precuneus, in the perceptual pro- cessing of familiar names, faces, and voices (von Kriegstein et al., 2005; Gorno-Tempini et al., 1998) and person-related semantic information (Tsukiura, Mochizuki-Kawai, & Fujii, 2006). We propose a role for the left STS/AG in accessing and integrating conceptual information related to target voices, in close communication with the regions planning and executing articulations. The increased performance demands encountered during the emulation of specific voice identities, which requires accessing the semantic knowledge of individuals, results in greater engagement of this left posterior temporo-parietal region and its en- hanced involvement with the speech production network. The interaction of right-lateralized sites on STS with left, middle, and inferior frontal gyrus and pre-SMA suggests higher-order roles in planning specific impersonations. Blank et al. (2002) found that the left pars opercularis of the IFG and left pre-SMA exhibited increased activation during production of speech of greater phonetic and lin- guistic complexity and variability and linked the pre-SMA to the selection and planning of articulations. In studies of voice perception, the typically right-dominant temporal voice areas in STS show stronger activation in response to vocal sounds of human men, women, and children com- pared with nonvocal sounds (Belin & Grosbras, 2010; Belin et al., 2000, 2002; Giraud et al., 2004), and right-hemisphere lesions are clinically associated with specific impair- ments in familiar voice recognition (Hailstone, Crutch, Vestergaard, Patterson, & Warren, 2010; Lang, Kneidl, l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j f . / t u s e r o n 1 7 M a y 2 0 2 1 Figure 5. Regions of the brain showing stronger positive correlations during the production of impersonations compared with accents. The seed regions were defined using from the contrast Impersonations >
Accents. Voxel height threshold p < .001 (uncorrected), cluster threshold p < .001 (corrected). 1882 Journal of Cognitive Neuroscience Volume 25, Number 11 Table 5. Brain Regions Showing an Enhanced Positive Correlation with Temporo-parietal Cortex during Impersonations, Compared with Accents Seed Region Right anterior STS Left posterior STS No. of Voxels Target Region 66 98 77 21 65 48 37 346 287 306 163 35 33 138 26 21 23 37 Left STG Right/left cerebellum Right cerebellum Left IFG (pars operc.) Right calcarine gyrus Left/right pre-SMA Right STG Left rolandic operculum/left STG/STS Left/right cerebellum Right STG/IFG Right/left caudate nucleus and right thalamus Left thalamus/hippocampus Left hippocampus Left pre/postcentral gyrus Left/right mid cingulate cortex Left IFG/STG Right postcentral gyrus Left insula/IFG Right posterior STS 225 Left middle/IFG 40 41 20 57 29 31 Left STS Right postcentral gyrus/precuneus Right IFG Left/right cerebellum Left lingual gyrus Left STG Coordinate x −60 9 15 −48 15 −3 63 −33 0 66 15 −12 −15 −51 −9 −57 54 −36 −39 −66 27 42 −24 −18 −63 y −12 −63 −36 9 −72 3 −33 −30 −48 −6 21 −27 −15 −6 9 12 −12 21 54 −36 −45 18 −48 −69 −6 z 6 −12 −18 12 18 51 9 18 −15 −3 3 −6 −21 30 39 3 36 3 0 6 57 27 −24 3 0 t 6.16 5.86 5.84 5.23 5.03 4.84 4.73 6.23 6.15 5.88 5.72 5.22 4.97 4.79 4.37 4.27 4.23 4.14 5.90 5.63 5.05 4.79 4.73 4.64 4.35 z 4.65 4.50 4.49 4.17 4.06 3.95 3.88 4.68 4.64 4.51 4.43 4.17 4.03 3.92 3.67 3.61 3.58 3.52 4.52 4.38 4.07 3.92 3.89 3.83 3.66 Voxel height threshold p < .001 (uncorrected), cluster threshold p < .001 (corrected). Coordinates indicate the position of the peak voxel from each significant cluster, in MNI stereotactic space. pars operc. = pars opercularis. Hielscher-Fastabend, & Heckmann, 2009; Neuner & Schweinberger, 2000). Investigations of familiarity and identity in voice perception have implicated both poste- rior and anterior portions of the right superior temporal lobe, including the temporal pole, in humans and ma- caques (von Kriegstein et al., 2005; Kriegstein & Giraud, 2004; Belin & Zatorre, 2003; Nakamura et al., 2001). We propose that the right STS performs acoustic imagery of target voice identities in the Impersonations condition, and that these representations are used on-line to guide the modified articulatory plans necessary to effect voice change via left-lateralized sites on the inferior and middle frontal gyri. Although there were some acoustic differences be- tween the speech produced under these two conditions— the Impersonations had a higher mean and standard deviation of pitch than the Accents (see Table 1)—we would expect to see sensitivity to these physical properties in earlier parts of the auditory processing stream, that is, STG rather than STS. Therefore, the current results offer the first demonstration that right temporal regions pre- viously implicated in the perceptual processing and rec- ognition of voices may play a direct role in modulating vocal identity in speech. The flexible control of the voice is a crucial element of the expression of identity. Here, we show that changing the characteristics of vocal expression, without changing the linguistic content of speech, primarily recruits left anterior insula and inferior frontal cortex. We propose that therapeutic McGettigan et al. 1883 D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j t / f . u s e r o n 1 7 M a y 2 0 2 1 approaches targeting metalinguistic aspects of speech pro- duction, such as melodic intonation therapy (Belin et al., 1996) and respiratory training, could be beneficial in cases of speech production deficits after injury to left frontal sites. Our finding that superior temporal regions previously identified with the perception of voices showed increased activation and greater positive connectivity with frontal speech planning sites during the emulation of specific vocal identities offers a novel demonstration of a selective role for these voice-processing sites in modulating the expression of vocal identity. Existing models of speech production focus on the execution of linguistic output and monitoring for errors in this process (Hickok, 2012; Price, Crinion, & Macsweeney, 2011; Tourville & Guenther, 2011). We suggest that noncanonical speech output need not always form an error—for example, the convergence on pronunciations observed in conversation facilitates comprehension, interaction, and social cohesion (Garrod & Pickering, 2004; Chartrand & Bargh, 1999). However, there likely exists some form of task-related error monitor- ing and correction when speakers attempt to modulate how they sound, possibly along a predictive coding mechanism that attempts to reduce the disparity between predicted and actual behavior (Price et al., 2011; Friston, 2010; Friston & Price, 2001)—this could take place in the right superior temporal cortex (although we note that previous studies directly investigating the detection of and compensation for pitch/time-shifted speech have located this to bilateral posterior STG; Takaso, Eisner, Wise, & Scott, 2010; Tourville et al., 2008). We propose to repeat the current experiment with professional voice artists who are expert at producing convincing impressions and pre- sumably also skilled in self-report on, for example, perfor- mance difficulty and accuracy. 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D o w n l o a d e d f r o m l l / / / / j t t f / i t . : / / h t t p : / D / o m w i n t o p a r d c e . d s f i r o l m v e h r c p h a d i i r r e . c c t . o m m / j e o d u c n o / c a n r a t r i t i c c l e e - p - d p d 2 f 5 / 1 2 1 5 / 1 1 8 1 7 / 5 1 1 8 9 7 4 5 6 / 0 1 7 4 7 7 o 9 c 6 n 8 _ 5 a / _ j 0 o 0 c 4 n 2 7 _ a p _ d 0 0 b 4 y 2 g 7 u . e p s t d o f n b 0 y 8 S M e I p T e m L i b b e r r a 2 r 0 2 i 3 e s / j . / t f u s e r o n 1 7 M a y 2 0 2 1 1886 Journal of Cognitive Neuroscience Volume 25, Number 11 Tʼainʼt What You Say, Itʼs the Way That You Say It—Left Insula image

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