Accepted Manuscript

Hemispheric Association and Dissociation of Voice and Speech Information

Processing in Stroke

Anna B. Jones, Andrew J. Farrall, Pascal Belin, Cyril R. Pernet, Dr

PII: S0010-9452(15)00245-2
DOI: 10.1016/j.cortex.2015.07.004
Reference: CORTEX 1531

To appear in: Cortex

Received Date: 9 January 2015

Revised Date: 22 May 2015
Accepted Date: 6 July 2015

Please cite this article as: Jones AB, Farrall AJ, Belin P, Pernet CR, Hemispheric Association and
Dissociation of Voice and Speech Information Processing in Stroke, CORTEX (2015), doi: 10.1016/
j.cortex.2015.07.004.

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 ACCEPTED MANUSCRIPT
Hemispheric Association and Dissociation of Voice and Speech
Information Processing in Stroke

Anna B. Jones1,2, Andrew J. Farrall1,2, Pascal Belin3,4 & Cyril R. Pernet1,2

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Brain Research Imaging Centre, The University of Edinburgh, UK
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Centre for Clinical Brain Sciences, The University of Edinburgh, UK

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Institute of Neuroscience and Psychology, University of Glasgow, United Kingdom
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Institut des Neurosciences de La Timone, UMR 7289, CNRS & Université Aix-Marseille, France

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Corresponding author:
Dr Cyril Pernet
Centre for Clinical Brain Sciences (CCBS)
The University of Edinburgh

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Chancellor's Building
Room GU426D
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49 Little France Crescent
Edinburgh EH16 4SB
[email protected]
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tel: 0131 465 9530

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Abstract

As we listen to someone speaking, we extract both linguistic and non-linguistic information.

Knowing how these two sets of information are processed in the brain is fundamental for the
general understanding of social communication, speech recognition and therapy of language
impairments. We investigated the pattern of performances in phoneme vs. gender categoriza-
tion in left and right hemisphere stroke patients, and found an anatomo-functional dissocia-
tion in the right frontal cortex, establishing a new syndrome in voice discrimination abilities.

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In addition, phoneme and gender performances were most often associated than dissociated
in the left hemisphere patients, suggesting a common neural underpinnings.

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Keywords

Voice gender perception, phoneme perception, hemispheric dissociation, stroke, aphasia

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Introduction

Speech perception is often seen as special (Liberman & Mattingly, 1989) because localized
brain injury can elicit specific language impairments such as aphasia, and because healthy
individuals are extremely efficient at categorizing phonemes and syllables despite large varia-
tions in the stimulus spectral patterns (Liberman, Delattre, & Cooper, 1952). To achieve high
performance levels, it has been hypothesized that voice information (talker specific infor-
mation) is extracted along with the speech signal, and then stripped away to access (invariant)
phonemic content: a process known as ‘speaker normalization’. This hypothesis is however

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challenged because general auditory learning mechanisms are capable of explaining category
formation in the absence of invariant acoustic information. Birds can learn speech consonant
categories with no obvious acoustic invariant cue (Kluender, Diehl, & Killeen, 1987) and

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human listeners can readily learn non-speech categories that are similarly structured (Wade &
Holt, 2005). In addition, several studies showed that talker variability influences speech per-
ception. For instance the literature describes increased memory for words spoken by familiar

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voices, compared to non-familiar voices (Nygaard & Pisoni, 1998; Nygaard, Sommers, &
Pisoni, 1994; Palmeri, Goldinger, & Pisoni, 1993), and similarly enhanced discrimination of,
and memory for, (non-familiar) speakers of our own language compared to speakers of an-
other language (Language Familiarity Effect - Perrachione & Wong, 2007) even in the ab-

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sence of intelligibility (Fleming, Giordano, Caldara, & Belin, 2014). Most of these studies do
not, however, specifically address the issue of phoneme perception, and thus acoustical regu-
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larities coming from multiple levels are at play.

Like speech perception, voice perception is often considered special (Belin, 2006; Belin, Fec-
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teau, & Bedard, 2004; Scott, 2008). Humans easily recognize different voices, and this ability
is of considerable social importance. Voice-selective areas have been demonstrated in the
human brain (Belin, Zattorre, & Ahad, 2002; Belin, Zattorre, Lafaille, Ahad, & Pike, 2000),
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localized bilaterally along the upper bank (middle and anterior) of the Superior Temporal
Sulcus (STS) (Alho et al., 2006; Belin, et al., 2002), and also in the inferior and orbitofrontal
cortex (Charest, Pernet, Latinus, Crabbe, & Belin, 2012; Fecteau, Armony, Joanete, & Belin
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2005) as well as surrounding insular cortex (Johnstone, van Reekum, Oakes, & Davidson,
2006; Rama et al., 2004). This neural selectivity for voice has also been established in other
mammals, in particular primates (Johnstone et al., 2006; Rama et al., 2004) and more recently
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dogs (Andics, Gacsi, Farago, Kis, & Miklosi, 2014). Given the presence of con-specific voice
neural selectivity in these animals, we can establish that neural voice selectivity is an old evo-
lutionary feature (~100 million years for a common ancestor between humans and dogs, ~25
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million years for a common ancestor between macaques and human), preceding the appear-
ance of speech (~5 to 2 million years ago for proto-language and ~150,000 to 300,000 years
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ago for speech - Perreault & Mathew, 2012). Following psycholinguistic studies suggesting
that phonetic attributes are an intrinsic component of the perception and identification of in-
dividuals, i.e. the recognition of voices (Remez, Fellowes, & Rubin, 1997), it is possible that
some brain regions dedicated to speech co-opted neurons already involved in con-specific
voice processing.

In this study, we investigated how phoneme and talker information processing relate to each
other, by comparing performances of right fronto-temporal (non-aphasic), left fronto-
temporal aphasic and left fronto-temporal non-aphasic stroke patients. Each participant cate-
gorized sounds from pitch equalized morphed continua as being male-female or /pa/-/ta/
(Pernet, Belin, & Jones, 2014). Stimuli were the same in both tasks, and participants had
therefore to discard talker specific or phoneme specific information depending on the task at

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hand. Given the importance of the right STS (Bestelemeyer, Belin, P., & Grosbras, 2011) and
right Middle and Inferior Frontal Gyrus (MFG-IFG) (Charest et al., 2012) in talker infor-
mation processing, we hypothesized that right hemisphere patients will show a dissociation
between the two tasks. In contrast, following our hypothesis of co-optation of voice selective
neurons in phoneme processing, we hypothesized that left hemispheric aphasic patients will
not show such dissociation, while non-aphasic patient could be impaired for voice but not
phoneme.

Materials and Methods

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The experiment used (program and stimuli) is freely available from
http://dx.doi.org/10.6084/m9.figshare.1287284. It runs under Matlab with the psychophysical

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toolbox (Brainard, 1997; Kleiner et al., 2007). The behavioral data and scripts used to analyze
the data are available from http://dx.doi.org/10.6084/m9.figshare.1287262. The imaging radi-
ological analysis is also available with the behavioral data. CT or MRI scans could not be

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shared as they belong to the UK National Health Servise (NHS) and not to the research team.
The study was approved by the NHS Lothian South East Scotland Research Ethics Commit-
tee 01 (REC reference number: 11/SS/0055) and NHS Lothian Research and Development
(R&D project number: 2011/W/NEU/09).

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Participants: Twenty-five stroke patients (14 males, 11 females) with a median age of 69
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years (min 39, max 85) were recruited into this study. At the time of testing, all patients were
at the chronic stage (median time between stroke and testing 90±17 days). Participants were
recruited as inpatients and outpatients from Lothian NHS hospitals via stroke physicians
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and Speech & Language Therapists between 10 and 60 weeks post-stroke with the sole inclu-
sion criterion of a stroke affecting perisylvian tissues (supplementary table 1). Exclusion cri-
teria were the presence of a previous stroke and/or English not being the participant’s first
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language.

All patients were tested for their mood (Visual Analogue Self Esteem Scale – VASES (Brum-
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fitt & Sheeran, 1999) and The Depression Intensity Scale Circles – DISCS (Turner-Stokes,
Kalmus, Hirani, & Clegg, 2005)) and language abilities (Western Aphasia Battery – WAB,
Shewan & Kertesz, 1980). No patient had language deficits in the group with right hemi-
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sphere lesions (N=9, 5 males and 4 females, WAB median score 98.8), and 10 out of 16 pa-
tients showed signs of aphasia in the left hemisphere group (N=10, 5 males and 5 females,
WAB median score 49.5 for aphasics vs. N=6, 4 males and 2 females, WAB median score
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99.4 for non-aphasics – percentile bootstrap difference 49.8 [15 80] p=0). Kruskall-Wallis
ANOVA showed that groups did not differ in terms of median age (χ2(2,22)=4.58 p=0.1), in
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median time delay between stroke and testing (χ2(2,22)=1.68 p=0.43) or depression scores
(χ2(2,22)=4 p=0.13 for VASES and χ2(2,22)=2.19 p=0.33 for DISCS). All demographic and
test results are summarized in supplementary table 2.

Paradigm: The experiment was identical to (Pernet et al., 2014), except that only pitch equal-
ized stimuli were used. Participants carried out with two 2 alternative forced choice identifi-
cation tasks: voice gender (male vs. female) and phoneme (/pa/ vs. /ta/), and responded by
button press on a keyboard. For each task, the same two continua of morphed sounds were
used: the 1st continuum going from a Male-/pa/ to a Female-/ta/ and the 2nd continuum with
the same speakers going from a Male-/ta/ to Female-/pa/. Morphs were generated by steps of
10% giving for the 1st continuum, 100% Male-/pa/, 90% Male-/pa/ with 10% Female-/ta/,
80% Male-/pa/ with 20% Female-/ta/, etc. until 100% Female-/ta/ and for the 2nd continuum,

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100% Male-/ta/, 90% Male-/ta/ with 10% Female-/pa/, 80% Male-/ta/ with 20% Female-/pa/,
etc. until 100% Female-/pa/. This design allowed investigation of the effect of the task while
controlling for the general acoustic characteristics of the stimuli, since the same stimuli were
used in both tasks. Participants heard each stimulus in pseudo-random order six times each,
for a total of 132 stimuli (2 continua * 11 steps * 6 trials) per task. Eighteen different contin-
ua of stimuli were generated from 6 different speakers (3 males and 3 females pronouncing
/pa/ and /ta/) and randomly assigned to participants. Task order and key orientation were
counterbalanced between participants. Between each task, an interfering tone discrimination
task was also performed. Participants heard pure tones of various frequencies corresponding

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to the male and the female ranges and had to tell if 2 consecutive sounds were the same or
different. The task followed a 2 down, 1 up step-wise procedure (Levitt, H., 1971) equating
participants performances when the staircase has ended (70.71% percent correct). This task

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was primarily designed to minimise the influence of one categorization task on the other, but
also allowed control for basic auditory impairments. No significant differences were observed
between groups on this task (Kruskall-Wallis ANOVA, χ2(2,22)=1.93 p=0.38, supplementary

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table 2).

Behavioral Classification: To assess the independence of phoneme and gender categorization

tasks, behavioral performances were binarized as impaired vs. unimpaired. For each subject,

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response proportion curves (percentage of female response or percentage of /ta/ responses)
were obtained by averaging repeated trials from the different continua (figure 1). Each partic-
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ipant was then classified based on his or her ability to perform outside chance level, at least
one time for the first 3 stimuli and at least one time for the last 3 stimuli along the sound con-
tinua. This implies that if a participant answered correctly for a least one of the initial stimuli
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(100% or 90% or 80% male or /pa/) and one of the final stimuli (100% or 90% or 80% fe-
male or /ta/), he or she was considered unimpaired (see supplementary material 1 for repeated
analyses using an incremental classification criterion). In normal heathy participants, this is
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achieved very easily. Taking the data from (Pernet et al., 2014), 100% of controls (N=18, 9
males, 9 females) were unimpaired (figure 2). From the resulting classification in patients, the
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independence between the phoneme and gender categorization performance was tested for
each group using a McNemar test with exact central probability (Fay, 2010).

Lesion to Symptom Mapping: Each participant received one or more CT or MRI scan from
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the National Health Service, and the scan the closest in time to the subjects’ participation in
this study was considered. One-to-one mapping was computed using a McNemar test with
exact central probability (Fay, 2010) between behavioral deficits (impaired/non-impaired)
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and 6 Regions Of Interest (lesioned/non-lesioned). Significance was considered at al-

pha=0.0083% i.e. Bonferroni corrected for the 6 ROI. The ROI classification was performed
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by an expert neuro-radiologist (AF) with 12 years of experience, and following a detailed

protocol1 adapted from the International Stroke Trial III (Whiteley et al., 2006). Regions of
Interest (ROI) considered were the middle/inferior frontal gyrus (involved in gender categori-
zation (Fecteau, et al., 2005, Charest et al., 2012), Heschl’s gyrus (also called Transverse
Temporal gyrus – involved in language learning and spectral/pitch information processing
(Warrier et al., 2009), the superior temporal gyrus (anterior/posterior, involved in general
auditory processes but also voice perception, Belin, et al., 2004), the insula (involved in cen-
tral auditory functions, in particular temporal resolution and sequencing (Bamiou et al., 2006)
and the amygdala-hippocampal complex (involved in memory and emotional voice percep-
tion (Johnstone et al., 2006; Rama et al., 2004).
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http://www.bric.ed.ac.uk/documents/CT_MR_for_voice_perception.pdf

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Behavioral quantitative analyses: Each participant ability to distinguish between male-female

and /pa/-/ta/ stimuli was investigated using signal detection theory (Macmillan, & Creelman,
2005), computing the perceptual distance d’ (i) between successive pairs of stimuli along
each continuum, and (ii) between the extreme stimuli of each continuum (global d’ - the dis-
tance between 100% male or /pa/ and 100% female or /ta/). We speak of categorical percep-
tion when a continuum of stimuli is divided by a boundary, such that stimuli on one side of
the boundary are perceived as belonging to one category, whereas stimuli on the other side
are perceived as belonging to another category, and perceptual distances within a category are

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low. Here we investigated categorical perception in each group and task by testing if d’ val-
ues differed from 0 (bootstrap-t test with Bonferroni correction - Wilcox, 2012) for each pair
of stimuli along the continuum, thus delineating a perceptual boundary. Finally, differences

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between gender and phoneme categorization performances were tested comparing global d’
values within groups (percentile bootstrap of the difference with adjustment for multiple test-
ing) and relative to the healthy participants from (Pernet et al., 2014) (mean difference be-

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tween groups, with adjustment for multiple comparisons based on the maximum statistics -
Wilcox, 2012).

Results

Behavioral classification
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Of the 25 patients recruited, 19 showed at least one categorization deficit (table 1), which we
defined as the inability to perform above chance for at least one of three extreme stimuli
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(100%, 90% or 80% male/pa, female/pa, male/ta or female/ta). As hypothesized, we observed

a significant dissociation in right fronto-temporal patients with impaired voice gender catego-
rization vs. intact phonological categorization (8 out of 9 patients, χ2=7, p=0, Φ=inf). In left
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fronto-temporal patients, we found no dissociation, for both the aphasic and the non-aphasic
groups. Aphasic patients tended to show both phonological and voice gender categorization
deficits (χ2=0.2, p=1, Φ=0.6) and non-aphasic patients tended to performed normally in both
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tasks (χ2=0, p=0.5, Φ=1 – figure 1). The same association/dissociation patterns were ob-
served when varying the categorization deficit criteria (supplementary material 1).
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[Table 1. Contingency table showing the classification of patients (per group) for the gender and the
phoneme tasks.]
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Lesion symptom mapping

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One-to-one mapping between the behavioral classifications (impaired / unimpaired) and re-
gion-of-interests (lesioned / not-lesioned) showed that gender categorization impairments are
associated with right frontal lesions (χ2=8, p= 0.0078 – see figure 2 for details). No other ROI
shows significant results.

[Figure 1. Average response proportion curves for each group of subjects (red lines represent pho-
neme task; blue lines represent gender task, bars represent bootstrap standard errors, shaded areas
indicate chance level from 21.09% to 78.91%). At the bottom is shown the distribution of lesions per
ROI for each group superimposed onto the MNI 152 template.]

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Behavioral quantitative analyses

Analyses of perceptual distances (d’) between successive pairs of items along continua re-
vealed that none of the patient groups had increased perceptual distances for ambiguous items
(figure 3), contrary to healthy subjects as shown in (Pernet et al., 2014). This result indicates
a generalized reduction in categorical boundaries following stroke. Further analyses on global
perceived distances (i.e. the d’ computed between 100% male and 100% female or 100% /pa/
and 100% /ta/ stimuli) showed that, compared to controls, right fronto-temporal patients had

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a lower global d’ in the gender categorization task only (difference controls vs. patients= 1.3,
adjusted 95% CI [0.8 1.8] for Male/Female, difference controls vs. patients= 0.06, adjusted
95% CI [-0.2 0.3] for Pa/Ta). For left fronto-temporal aphasic patients, global d’ was lower

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than in controls for both male/female stimuli (difference controls vs. patients= 1.4 adjusted
95% CI [0.5 2.2]) and pa/ta stimuli (difference controls vs. patients= 1.4, adjusted 95% CI
[0.4 2.4]). For left fronto-temporal non-aphasic patients, global d’ did not differ from controls

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(difference controls vs. patients= 0.11 adjusted 95% [-0.27 0.49] for Male/Female, difference
controls vs. patients= 0.17, adjusted 95% CI [-0.49 0.83] for Pa/Ta). Finally, looking at with-
in group differences, whilst right fronto-temporal patients show a significant difference be-
tween tasks, no significant differences were observed for both left fronto-temporal groups

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(figure 1, table 2), which concurs with classification results.
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[Table 2. Global perceived distance (d’) computed for each group with within group differences, ad-
justed 95% confidence intervals and p values.]
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[Figure 2. At the top left is shown the response proportion curves in normal healthy subjects (Pernet
et al., 2014), illustrating that the classification works for well with controls scoring at 100% correct.
Below are the same response proportion curves but for each right hemisphere patient, associated with
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axial slices showing the lesion in relation to the Sylvian fissure (highlighted in yellow). Importantly,
the STG is intact in 7 out of 9 cases (patients 3 and 5 having both frontal and temporal lesions), sug-
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gesting that this region is not critical for gender/voice categorization.]

[Figure 3. Perceptual and global perceptual distances (d’). Curves show d’ values in the gender cat-
egorization task (blue) and the phoneme categorization task (red). Bars represent bootstrap standard
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errors. Box plots show the global d’ and the difference in global d’ between the two tasks for control
healthy participants (HP - black), left aphasic (LA - green), left non-aphasic (LNA - red) and right
non-aphasic (RNA - blue) patients. Boxes represent the interquartile range with the median, and
whiskers show the 1.5*interquartile distance with outliers marked beyond that points.]
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Discussion
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Based on functional MRI (Belin, et al., 2002, 2000; Charest et al., 2012) and Transcranial
Magnetic Stimulation (Bestelemeyer, et al., 2011) results observed in healthy volunteers, we
hypothesized that patients with right fronto-temporal stroke will show a deficit in gender cat-
egorization but intact phonological performances. Our results using both a qualitative (classi-
fication of percentage of responses) and a quantitative (signal detection theory) approach con-
firmed this hypothesis. To our knowledge this is the first time that such deficit has been de-
scribed in the literature. All of the patients presenting right frontal lesions showed a deficit in
voice categorization, thus demonstrating a significant brain/behavior association. Previous
studies have reported cases of phonagnosia (Van Lancker & Canter, 1982; Van Lancker,
Cummings, Kreiman, & Dobkin, 1988; Van Lancker, Kreiman, & Cummings, 1989), in

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which patients could not recognize familiar voices; but this deficit was associated with right
parietal lesions. When discrimination of unfamiliar voices was tested, deficits were associat-
ed with temporal (left or right) lesions although some evidence also exists for voice deficit
during fronto-temporal degeneration (Hailstone, Crutch, Vestergaard, Patterson, & Warren,
2010; Hailstone, Crutch, & Warren, 2010). What remains unclear is (1) if the deficit is specif-
ic to gender categorization or if it also relates to identity and (2) what is the role of the right
IFG. On one hand, studies comparing attention to voice vs. speech found voice specific ef-
fects over the right STS (von Kriegstein, Eger, Kleinschmidt, & Giraud, 2003), and this effect
has been related to speaker identity (Schall, Kiebel, Maess, & von Kriegstein, 2014). On the

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other hand, using a continuous carry over-over design allowing acoustic from perceived dis-
tances to be distinguished during gender categorization, Charest et al., (2012) showed that the
STS processes gender (and thus identity as well) related acoustic information whilst the right

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IFG is involved in perceived gender related distances. In most studies, stimuli are pitch equal-
ized. We previously showed that equalizing pitch does not influence performance or RT
(Pernet & Belin, 2012; Pernet et al., 2014); thus making timbre and consequently spectro-

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temporal analysis a key element in gender categorization. There is no doubt that the patients
in our study found the gender categorization task difficult, even when the STS was intact
(figure 2 – patients 1, 2, 4, 6, 7, 8, 9). This difficulty was not related to the absence of pitch
information since their pitch perception threshold did not differ from other patient groups. It

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has been proposed that the right IFG plays a general role in voice recognition and social
communication at large, since direct connections have been demonstrated between the STG
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and the ventrolateral prefrontal cortex of the macaque (equivalent of IFG in humans) along
with vocalization responsive cells (Romanski, 2012). Voice deficits observed here following
right IFG lesions, often in association with insula lesions (although the association was not
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significant), could then reflect a disconnection syndrome from the STS.

Results observed in left hemisphere patients were heterogeneous. The absence of one-to-one
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mapping between speech deficits and brain lesions is not completely surprising since many
studies have found that phonological categorization depends critically on the left supra-
marginal gyrus, a region not investigated in our study. Phonological deficits were also not
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always associated with aphasia (4 out of 16 patients), which concurs with the idea that sub-
lexical speech perception impairments do not necessarily predict auditory comprehension
deficits (Turkeltaub & Coslett, 2010). More than half of the left hemisphere patients (9 out of
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16) were either impaired or unimpaired in both tasks, and in the aphasic group 4 out of 10
patients showed a double deficit and 5 others showed reduced performances, resulting in
lower perceived distances between extreme stimuli (either male-female or /pa/-/ta/). One pos-
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sible explanation is that aphasic patients simply did not understand the instructions. Because
these same patients could, however, perform other tasks with more complex instructions (e.g.
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the tone discrimination task used to check their pitch perception - see method), we consider
this pattern of results as supportive of the co-optation hypothesis. Indeed, if we accept that
there is speech-selectivity in the left fronto-temporal cortex, then we have to conceive that
this selectivity can be associated with the mechanisms that produce and perceive the sounds
of speech (McGettigan & Scott, 2012). The dissociations observed in 3 left hemisphere pa-
tients, also reveals that gender categorization deficits can be observed following left hemi-
sphere lesions, and therefore that gender categorization is processed bilaterally.

In conclusion, through the analysis of categorization performances of right fronto-temporal

stroke patients, we showed that the right frontal cortex (likely the ventral part of the IFG)
plays a major role in voice/gender information processing. In contrast, left fronto-temporal
patients (aphasic or not) tend to show associated performances for both voice/gender percep-

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tion and speech perception, although dissociation are also possible. Together, these results
lend support to the hypothesis of bilateral processing of voice information with (i) a im-
portant role of the right frontal cortex in voice categorization and (ii) both common and dedi-
cated mechanisms, in the left hemisphere, for talker and speech information processing.

Acknowledgment

This work was supported by SINAPSE (http://sinapse.ac.uk/) who funded A.B. Jones.

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Gender task
Unimpaired Impaired
Left fronto- Unimpaired 1 2
temporal
Impaired 3 4
Aphasics
Left fronto- Unimpaired 4 1
Phoneme task
temporal Non-
Impaired 1 0
aphasics

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Right fronto- Unimpaired 1 7
temporal Impaired 0 1
Table 1. Contingency table showing the classification of patients (per group) for the gender and the

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phoneme tasks.

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Left fronto- Left fronto-
Right fronto-
Controls temporal temporal Non
temporal
Aphasics Aphasics
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Male/Female
Distance Pa/Ta 2.55 1.1 2.17 2.49
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Difference and 0.01 [-0.1 0.1] 0.019 [0.9 0.8] 0.054 [-0.4 0.6] 1.25 [1.8 0.7]
adjusted 95% CI p=0.872 p=0.958 p=0.86 p=0
Table 2. Global perceived distance (d’) computed for each group with within group differences,
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adjusted 95% confidence intervals and p values.

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Subject Stroke type Arterial Regions of interest showing lesion WAB AQ and
territory classification
1 Ischemic Left MCA 'IFG' 'insula' 'anterior STS' 'posterior 19 -Global
1,2,3,4,5,6 STS' 'Heshl gyrus'
2 Ischemic Left MCA 'IFG' 'insula' 'Heshl gyrus' 38.15 -Transcortical
1,2,4,5 Motor
3 Ischemic Left MCA 1,4 'IFG' ‘MFG’ 60.9 - Broca's
4 Ischemic Left MCA 4,6 'IFG' ‘MFG’ 89.6 - Anomia
5 Hemorrhagic Left MCA 'IFG' 'anterior STS' 'posterior STS' 17 - Wernicke's

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1,2,3,6
6 Ischemic Left MCA 1 'IFG' 'insula' 'Heshl gyrus' 36.6 - Broca's
7 Ischemic Left MCA 1 'IFG' 'insula' 20.8 – Broca’s

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8 Ischemic Left MCA 2, 'IFG' 'insula' 'anterior STS' 'posterior 62.9 - Wernicke's
3,6 STS' 'Heshl gyrus'
9 Ischemic Left ACA 'IFG' 83.6 - Anomia

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10 Ischemic Left MCA 'IFG' 'insula' 'anterior STS' 'posterior 65.1 - Conduction
2,3,6 STS' 'Heshl gyrus'
11 Ischemic Left MCA 5 'IFG' 99.4
12 Ischemic Left LSA 'IFG' 'insula' 97.9

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13 Ischemic Left PCA 'insula' 'Amygdala-Hippocampal 99.4
complex'
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14 Ischemic Left MCA 'IFG' 100
4,5,6
15 Ischemic Left MCA 1 'IFG' 'insula' 'Heshl gyrus' 99.4
16 Ischemic Left LSA 'insula' 100
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17 Ischemic Right MCA 'IFG' 98.4

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18 Hemorrhagic Right MCA 'IFG' 'insula' 98.8
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2,4,5 STS' 'Heshl gyrus'
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20 Ischemic Right LSA 'IFG' 98.8

21 Ischemic Right MCA 2, 'IFG' 'insula' 'anterior STS' 'posterior 100
5 STS' 'Heshl gyrus'
22 Hemorrhagic Right MCA 5 'IFG' 100
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23 Ischemic Right MCA 1 'IFG' 'insula' 98.6

24 Ischemic Right MCA 'IFG' 98.8
1,2, 3, 4, 5, 6
25 Ischemic Right MCA 1 'IFG' 'insula' 93
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Supplementary table 1. Patient clinical scan evaluations based on the International Stroke Trial III
(Whiteley, Lindsey, Wardlaw, & Sandercock, 2006) along with the aphasia quotient and classification
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(Shewan & Kertesz, 1980). PCA stands for posterior cerebral artery. MCA stands for middle cerebral
artery, and 1 to 6 refers to sub-territories (1 small cortical, 2 basal ganglia, 3 lateral to the ventricle,
4 anterior half of peripheral MCA, 5 posterior half of peripheral MCA, 6 whole peripheral MCA).
ACA stands for anterior cerebral artery. LSA stands for lenticulo-striate arteries. IFG: inferior
frontal gyrus, STS superior temporal sulcus.
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Left Aphasic Left Non Aphasic Right Non Aphasic

Sample size 10 6 9
Age (years) 71 ± 3.8 61 ± 8.34 65 ± 4.7
Stroke to Test time (days) 89 ± 18 133 ± 83 86 ± 58
Tone discrimination
8.25 ± 1.16 5.37 ± 2.31 6.75 ± 1.09
threshold (Hz)
WAB score (out of 100) 49.5 ± 14 99.4 ± 0.5 98.8 ± 0.3
VASES (out of 50) 35 ± 3.4 44.5 ± 2.9 37 ± 4.9
DICS (out of 4) 2.5 ± 0.4 2 ± 0.39 3±1

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Supplementary table 2. Medians with standard errors of patient demographics, basic auditory,
language, and depression characteristics.

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Supplementary material 1

Changing the classification criterion to the inability to perform above chance for extreme
stimuli only, then for at least one of two, at least one of three, at least one of four and at least
one of five of extreme stimuli, gave very similar results with 20/25, 19/25, 19/25, 18/25 and
18/25 of patients classified impaired. Importantly, the number of patients showing
dissociations was relatively stable, such as associations/dissociations results were unchanged
by changing the criterion (see supplementary figure 1). In the left aphasic group, out of the 10
patients, the number of subjects showing dissociations was respectively 4, 4, 5, 5 and 5. In

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the left non aphasic group, the same 2 patients always showed a dissociation. Finally in the
right hemisphere group, out of 9 patients, the number of subjects showing dissociations was
respectively 7, 7, 7, 6 and 6.

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[Supplementary figure 1. Percentage of patients impaired in each task for each patient group,
computed using multiple criteria (red lines represent phoneme task; blue lines represent gender task,

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vertical bars represent the 95% confidence interval).]

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