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Mécanismes neurophysiologiques de la perception de

flux sonores chez l’Homme : effets des contexts


acoustiques et attentionnels.
Aurélie Bidet-Caulet

To cite this version:


Aurélie Bidet-Caulet. Mécanismes neurophysiologiques de la perception de flux sonores chez l’Homme :
effets des contexts acoustiques et attentionnels.. Neurosciences [q-bio.NC]. Université Claude Bernard
- Lyon I, 2007. Français. �tel-00189055�

HAL Id: tel-00189055


https://tel.archives-ouvertes.fr/tel-00189055
Submitted on 19 Nov 2007

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UNIVERSITE CLAUDE BERNARD - LYON I

Président de l’Université M. le Professeur L. COLLET


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Attention, MCT

Reconnaissance de la source sonore

Traitement de la structure temporelle


Extraction des attributs auditifs

Groupement

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( retrait partie des lobes frontaux et )
Cortex c r bral

Cortex auditif secondaire

Cortex auditif primaire


Dienc phale

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Dynamics of a Temporo-Fronto-Parietal
Network during Sustained Spatial
or Spectral Auditory Processing

Aurélie Bidet-Caulet and Olivier Bertrand

Abstract
& Animal and human studies have suggested that posterior a temporo-parieto-frontal network was activated during the
temporal, parietal, and frontal regions are specifically involved whole duration of the stream in all conditions and modulated
in auditory spatial (location and motion) processing, forming a by attention; (ii) the left superior temporal cortex was the only
putative dorsal ‘‘where’’ pathway. We used scalp EEG and cur- region showing different activations for pitch and spatial varia-
rent density mapping to investigate the dynamics of this net- tions. Therefore, parietal and frontal regions would be involved
work in human subjects presented with a varying acoustic in task-related processes (attention and motor preparation),
stream in a two-factor paradigm: spatial versus pitch variations, whereas the differential processing of acoustic spatial and object-
focused versus diverted attention. The main findings were: (i) related features seems to take place at the temporal level. &

INTRODUCTION
Bates, & Goldman-Rakic, 1999; Romanski, Tian, et al.,
Both neurophysiological works in animals and human 1999). By analogy with the visual system, these ventral/
brain imaging studies have suggested separate auditory dorsal routes were suggested to support two function-
processing pathways for object-related and spatial fea- ally distinct ‘‘what’’ and ‘‘where’’ auditory pathways
tures, similarly to the visual system (Rauschecker, 1998). (Kaas & Hackett, 1999).
Within the lateral belt of the auditory cortex of anaes- In humans, many lesion (Clarke et al., 2002; Griffiths,
thetized macaque monkeys, neurons with narrow spatial Rees, Witton, et al., 1996), functional neuroimaging
tuning were situated in the caudal region, whereas a (Hart et al., 2004; Rama et al., 2004; Pavani, Macaluso,
better selectivity to monkey vocalizations was found in Warren, Driver, & Griffiths, 2002; Warren, Zielinski,
the rostral region (Tian, Reser, Durham, Kustov, & Green, Rauschecker, & Griffiths, 2002; Zatorre, Bouffard,
Rauschecker, 2001; Rauschecker & Tian, 2000). Two Ahad, & Belin, 2002; Alain, Arnott, Hevenor, Graham, &
recent functional MRI studies reported the same func- Grady, 2001; Maeder et al., 2001; Griffiths, Green, Rees,
tional dissociation within the temporal auditory cortex & Rees, 2000; Lewis, Beauchamp, & DeYoe, 2000;
in humans: anterior lateral auditory areas (lateral Baumgart, Gaschler-Markefski, Woldorff, Heinze, &
Heschl’s gyrus, anterior planum temporale, and planum Scheich, 1999; Bushara et al., 1999; Griffiths & Green,
polare) are specifically activated by pitch changes, 1999; Weeks et al., 1999; Griffiths, Rees, Rees, et al.,
whereas posterior medial auditory regions (posterome- 1998), and electrophysiological (Kaiser & Bertrand,
dial planum temporale) are activated by spatial location 2003; Kaiser, Ripper, Birbaumer, & Lutzenberger, 2003;
changes (Hart, Palmer, & Hall, 2004; Warren & Griffiths, Ducommun et al., 2002; Lutzenberger, Ripper, Busse,
2003). Connectivity studies in macaque have identified Birbaumer, & Kaiser, 2002; Xiang et al., 2002; Alain
(i) a ventral route connecting rostral auditory belt and et al., 2001; Anourova et al., 2001) data are consistent
parabelt areas, rostral superior temporal gyrus and ven- with the hypothesis of ‘‘what’’ and ‘‘where’’ separate
tral prefrontal cortex, and (ii) a dorsal route connecting auditory streams: anterior temporal and inferior frontal
caudal auditory belt and parabelt areas, and parietal and regions would process auditory ‘‘object’’ information
dorsolateral prefrontal cortices (Kaas & Hackett, 2000; and posterior temporal areas, parietal and frontal cor-
Rauschecker & Tian, 2000; Hackett, Stepniewska, & tices are involved in the spatial processing of both
Kaas, 1999; Kaas, Hackett, & Tramo, 1999; Romanski, stationary and moving sounds. This dual pathway was
recently supported in a neuroimaging meta-analysis by
Arnott et al. (2004).
INSERM U280, Mental Processes and Brain Activation Labora- However, the extent and the functional role of these
tory, IFNL, UCBL1, Lyon, France two pathways remain unclear and controversial (Hall,

D 2005 Massachusetts Institute of Technology Journal of Cognitive Neuroscience 17:11, pp. 1691–1703
!!     

2003; Middlebrooks, 2002; Belin & Zatorre, 2000; Cohen tively, p > .05), or within the attention task, between
& Wessinger, 1999). Indeed, the parietal areas do not spatial and pitch variation processing (93.2% and 92.9%,
seem to be specifically involved in auditory spatial respectively, p > .05). All subjects reported that they
processing and could also be activated during pitch- were mentally following spatial or pitch variations of the
oriented tasks (Maeder et al., 2001; Rama et al., 2000; auditory streams in the AS and AP conditions while they
Weeks et al., 1999; Zatorre, Mondor, & Evans, 1999; were actively ignoring the auditory stream trajectories in
Zatorre, Evans, & Meyer, 1994). the control task. Moreover, the EOG signal was drifting
In a recent neuroimaging study (Zatorre, Bouffard, during stimulus presentation and no attention or acous-
et al., 2002), a clear parietal activation could be found tic variation effect was noticed in the EOG time courses
only when transforming spatial auditory information ( p > .05) until 700 msec after variation onset.
into an oriented joystick movement. Moreover, the
activation of the parieto-frontal network, during audio-
spatial processing, could be due to goal-directed
Topographical Analysis
selective attention in active paradigms, as well as
stimulus-driven attentional capture (Behrmann, Geng, The main component elicited by the auditory stimulus
& Shomstein, 2004). In passive listening paradigms, an onset, N1, showed a classical ERP topography with a
involuntary attentional switch towards more attractive fronto-central maximum and a polarity reversal across
stimuli, such as moving sounds for instance, could the sylvian fissure, in both A and C conditions (Figure 1).
account for parietal activities (Pavani et al., 2002; Warren Computing scalp current densities (SCDs) by surface
et al., 2002; Xiang et al., 2002; Griffiths, Green, et al., laplacian has proved to be efficient in separating tem-
2000; Griffiths & Green, 1999; Griffiths, Rees, Rees, et al., poral, frontal (Giard et al., 1994), and parietal (Kaiser &
1998). Thus, the dorsal auditory pathway, usually con- Bertrand, 2003) components of auditory responses.
sidered as specifically processing spatial auditory infor- Applied to the present data, SCD maps showed, in
mation, could rather be involved in nonstimulus-specific both tasks, bilateral current sink/source patterns over
cognitive processes. temporal regions and a current sink over fronto-central
The present study aims at clarifying the role of the areas (Figure 1). These components were maintained
different components of the temporo-parieto-frontal throughout the stimulus duration.
network during the processing of pitch-related or spatial After this N1 component, we found a parieto-occipital
acoustic features. We used ERP mapping in human component largely prominent in the attention condition
subjects processing varying acoustic streams in situa- (Figure 2). SCD analysis permitted to dissociate two
tions of controlled attention. By means of a two-factor subcomponents: a current source over superior parietal
paradigm (spatial vs. pitch variations, focused vs. diverted cortices and bilateral sources over temporo-parieto-
attention), we could characterize brain regions specifi- occipital (TPO) junctions, clearly visible on the
cally involved in the processing of pitch and/or spatial attention-minus-control maps (Figure 2).
acoustic features and investigate how their activity could In order to estimate the time courses of those com-
be modulated by attention. Furthermore, by analyzing ponents, SCD maps were averaged across subjects and
the dynamics of the temporo-parieto-frontal network, it subsets of electrodes were selected to cover eight
became possible to dissociate components related to regions. Fronto-central (F: Fz/Cz), right/left superior
the sensory integration of specific acoustic features from temporal (RST: C4 and LST: C3), and right/left inferior
those components corresponding to higher-level cogni- temporal (RIT: T8/P8 and LIT: T7/P7) electrode groups
tive processes, such as attention for instance. were defined from the N1 topography (Figure 1),
whereas superior parietal (P: CP1/Pz/CP2) and right/left
temporo-parieto-occipital (RTPO: PO4 and LTPO: PO3)
electrode groups were from the topographies of the
RESULTS
varying period (Figure 2). SCD curves were averaged
Two participants had to be excluded because of exces- within each group for subsequent statistical analysis
sive eye blinks or muscular activity. Thus, 14 subjects (Figures 3 and 4).
were included in the analysis considering four condi-
tions: attention oriented (A) to the streams with spatial
(AS condition) or pitch (AP) variations, and attention
Fronto-Central Component
diverted (control, C) from acoustic feature variations
(CS and CP, respectively). The frontal component (F in Figure 3) started with a
transient N1/P2 complex followed by a sustained nega-
tive def lection throughout the stimulus duration
Behavioral Data
(emerging at 210 msec for all conditions, p < .001), on
Correct response rates did not differ between attention which a small N1/P2 complex was superimposed at
and control tasks (mean value, 93.1% and 92.2%, respec- variation onset. During the stationary period, both

1692 Journal of Cognitive Neuroscience Volume 17, Number 11


?   !           

for the P2 component. During the varying part, no


attention, acoustic variation, or interaction effect was
found significant for the sustained component.

Temporal Components
The temporal N1/P2 complexes were followed by a
sustained bilateral def lection with polarity reversal,
maintained during the entire stream duration (emerging
at 210 msec for all conditions, p < .01 at RST, p < .05 at
LST, and p < .01 at LIT).
During the stationary period, N1 mean amplitude (80–
130 msec) to auditory stream onset was significantly
more negative for the attention than the control condi-
tion at LST ( p < .05), RST ( p < .05), and LIT ( p < .001)
sites. The sustained waves were partially enhanced in
Figure 1. Stream-onset N1 topographies. Left hemisphere ERP the A condition during the stationary period as indicated
and SCD maps (mean value between 80 and 130 msec, 14 subjects)
in attention and control conditions. SCD allows to better disclose
by gray bars in Figure 3. No effect was detected for the
temporal and frontal components. Black ovals indicate electrode P2 component.
groups chosen for the analysis: fronto-central current sink (F), left At stream variation onset, a small N1/P2 complex was
superior temporal current sink (LST), and left inferior temporal superimposed on the sustained waves. During the vary-
current source (LIT). Very similar topographies were obtained ing period, pitch variations induced a larger sustained
over the right hemisphere and symmetrical electrode groups were
chosen for the right temporal components.
activity than spatial variations over the left auditory
cortex, irrespective of attention (LST: all 210-msec
windows from 385 to 910 msec, p < .01 and LIT: all
stream-onset N1 mean amplitude (80–130 msec) and the 210-msec windows from 595 to 910 msec, p < .05). Ad-
sustained negative deflection (210–420 msec, p < .05, ditionally, an interaction effect (Attention  Acoustic
and 525–735 msec, p < .05) were more negative for the variations) was found at the LST site (all 210-msec
attention condition ( p < .001). No effect was detected windows from 490 to 910 msec, p < .05). Wilcoxon tests

Figure 2. Topographies during the varying period. ERP and SCD maps (left, back, and right views, mean value between 385 and 595 msec
after variation onset, 14 subjects) in attention and control conditions, and difference maps between conditions. In ERP maps, a positive component
is present in the attention condition and is topographically distributed over parietal and occipital areas. SCD maps allow to disclose distinct
and focal current sources over the superior parietal cortex for both tasks and over TPO junctions in the attention condition only, clearly visible
on the attention-minus-control maps. Ovals indicate the electrode groups chosen for the analysis: superior parietal current source (P), right
and left TPO current sources (RTPO and LTPO, respectively), in addition to the frontal and temporal components already defined in the N1
topography (Figure 1).

Bidet-Caulet and Bertrand 1693


!!     G

Figure 3. Temporal and frontal SCD time courses. Mean SCD curves (14 subjects) are drawn over the three analysis periods (defined in
Figure 6C) for the frontal (F ), left superior and inferior temporal (LST and LIT, respectively), and right superior and inferior temporal (RST
and RIT, respectively) electrode groups. The time structure of the stimulus is shown at the top of the figure (gray ovals represent additional
noise-bursts used in the control task). During the stationary part, only mean attention (A) and control (C) curves are drawn, whereas in the
varying part all conditions are depicted with different line symbols. The evoked N1/P2 complex to auditory stream onset is clearly visible at all
electrode groups. N1/P2 to acoustic variations onset can also be noticed, although less pronounced. Note the polarity reversal throughout the
stimulus duration between superior and inferior temporal components bilaterally. Significant differences between the attention and the control
conditions are indicated by gray bars (ANOVA: *p < .05, **p < .01, ***p < .001). During the varying part, the left temporal components show
a significantly more pronounced sustained response for pitch (AP and CP) than for spatial variations (AS and CS) with stars symbolizing the
level of significance (ANOVA: *p < .05, ***p < .001). The black horizontal bars at the LST site indicate interaction effects between attention
and acoustic variation ( p < .05).

1694 Journal of Cognitive Neuroscience Volume 17, Number 11


   !           

Figure 4. Parietal SCD time courses. Mean SCD curves (14 subjects) are presented for the three analysis periods (defined in Figure 6C) for
the LTPO and RTPO, respectively, and the superior parietal (P) electrode groups. The time structure of the stimulus is shown at the top of the
figure (gray ovals represent additional noise-bursts used in the control task). During the stationary part, only mean attention (A) and control
(C) curves are drawn, whereas in the varying part all conditions are depicted with different line symbols. Note the large TPO activities developing
in the attention condition only and a sustained parietal activity enhanced by attention. Significant differences between the attention and the
control conditions are indicated by gray bars (ANOVA: *p < .05, **p < .01, ***p < .001).

showed that the negativity was significantly larger for the sponse (35 msec) elicited by the amplitude modulation
AP condition than for AS and CP ( p < .01), and for CP of the auditory stream.
than CS ( p < .05), whereas no significant difference was
found between the AS and CS conditions. The same
Temporo-Parieto-Occipital Components
effects were found during the final period as indicated
in Figure 3. Bilateral positive sustained components (LTPO and
The apparent ‘‘noise’’ on these temporal waveforms RTPO in Figure 4) were significantly emerging, over the
actually corresponded to the periodic steady-state re- TPO junctions, in the A condition only, from 250 msec

Bidet-Caulet and Bertrand 1695


!!     

to the end of the stationary period ( p < .05) and during both the stationary and the varying parts in the
from 350 msec after the variation onset until the end attention condition only; and (iii) a left temporal com-
of the stream ( p < .05). In the A condition, irrespective ponent differentially activated by spatial and pitch acous-
of the type of acoustic variations, these components tic variations.
were significantly more positive during the stationary
(all 210-msec windows from 210 to 735 msec, p < .05),
Attention Network
varying (all 210-msec windows from 280 to 910 msec,
p < .01), and final (all 210-msec windows from 840 to As parietal and frontal activities were maintained during
420 msec, p < .01) periods. No difference between the whole duration of the stream and were only mod-
pitch and spatial variations, no interaction effect, and no ulated by the orientation of attention and not by the
interhemispheric difference were found significant for acoustic feature to be integrated, they are likely to be
these components. related to high-level task-related processes.

Superior Parietal Component Fronto-Central Component


A positive sustained component (P in Figure 4) was The sustained fronto-central negative component was
significantly emerging, over superior parietal cortices, enhanced in the attention task during the stationary
in all conditions, from 100 msec to the end of the stream part. This could reflect the subject’s expectation of the
( p < .05). This component was significantly more next critical event (i.e., the acoustic variation onset).
positive in the A condition during the stationary (all During the varying part, this component did not differ
210-msec windows from 210 to 735 msec, p < .01), between conditions and could be associated with com-
varying (all 210-msec windows from 280 to 910 msec, p < mon processes required by both tasks: the anticipatory
.001), and final (all 210-msec windows from 840 to attention to the imperative acoustic event (end of the
420 msec, p < .001) periods. No difference between stream or noise bursts) and the preparation of the
pitch and spatial variations and no interaction effect was motor response. It was similar, in topography and time
found significant for this component. There was no course, to the frontal component reported in a previous
significant interhemispheric difference (Wilcoxon test, working memory study during the expectation of the
p > .05) when comparing SCD activities at the left and crucial stimuli (Kaiser & Bertrand, 2003), and could be
right parietal electrodes (Cp1 and Cp2). considered as a contingent negative variation compo-
nent (Brunia & van Boxtel, 2001).

DISCUSSION Superior Parietal Component


ERP correlates of spatial or pitch variation processing The sustained superior parietal activity was emerging in
were investigated during a highly demanding auditory both tasks and enhanced in the attention condition
task requiring the continuous processing of stream during the whole duration of the stream. This activity
trajectory, contrasted to a similarly difficult control could be due to eye movement planning (Andersen,
condition with diverted attention. Using similarly com- 1995), because we found slow EOG drifts during stimu-
plex pitch and spatial trajectories enabled to charac- lus presentation with no difference between tasks dur-
terize the brain regions involved in such feature ing the varying part.
integration. SCD mapping permitted to both disen- Parietal subregions have also been shown to be
tangle several active brain structures and estimate their involved in ‘‘multisensory integration and coordinate
dynamics. It then became possible to dissociate compo- transformation required to convert sensory input to
nents related (i) to basic auditory integration (sustained motor output’’ (Andersen & Buneo, 2002). In the pres-
during the whole stream duration), (ii) to sensory ent study, subjects had to achieve a complex transfor-
integration of specific acoustic features (during the mation converting the acoustic input into an oriented
varying period only), (iii) to cognitive processes re- joystick movement (right–left or front–back). That could
quired by the tasks and common to both features explain why both tasks activate the parietal cortex. This
(sustained during the whole stream duration). Further- interpretation is consistent with recent neuroimaging
more, by comparing two active tasks with identical results (Zatorre, Bouffard, et al., 2002) showing a clear
stimuli and similar demands on arousal and motor parietal activation only when transforming spatial audi-
response preparation, we could investigate how atten- tory information into a directed joystick movement.
tion modulated these activities. The most important Finally, in the present study, the superior parietal
findings were: (i) a fronto-parietal network activated activity could simply reflect supramodal attentional pro-
during the whole duration of the stream in all conditions cesses (Andersen & Buneo, 2002). In the attention
and modulated by attention, irrespective of acoustic condition, the parietal activity enhancement could
features; (ii) a sustained activity over the TPO junctions be due to selective attention, including the voluntary

1696 Journal of Cognitive Neuroscience Volume 17, Number 11


   !           

allocation of attention to spatial or pitch features of the


auditory stream (Behrmann et al., 2004; Zatorre, Mon-
dor, et al., 1999; Pugh et al., 1996). Interestingly, in a
recent study, a superior parietal cortex activation has
been found for attention to sound motion irrespective
of the stimulus properties (spectro-temporal or spatial)
(Hart et al., 2004). In addition, Warren and Griffiths
(2003), in a passive paradigm of spatial versus pitch
auditory perception, did not find any parietal activity
and explained it by ‘‘the lack of an output task.’’ Thus,
the parietal areas, which were proposed as contributing
to auditory spatial processing by several authors (Hart
et al., 2004; Zatorre, Bouffard, et al., 2002; Alain et al.,
2001; Maeder et al., 2001; Lewis et al., 2000; Bushara
et al., 1999; Weeks et al., 1999), may as well be involved
in task-related processes such as auditory attention and/ Figure 5. Surface electrode positions on a normalized brain with
or motor preparation. Besides, in passive listening par- neuroimaging findings. The Talairach coordinates of activations
adigm (Pavani et al., 2002; Warren et al., 2002; Griffiths, induced by visual motion in area V5 (diamonds) were collected in
Green, et al., 2000; Griffiths & Green, 1999; Griffiths, several neuroimaging studies (Dumoulin et al., 2000; Lewis et al.,
2000; Sunaert, Van Hecke, Marchal, & Orban, 1999, 2000; Chawla
Rees, Rees, et al., 1998), these parietal activities could
et al., 1999; Goebel et al., 1998). These activations were superimposed
be due to stimulus-driven attentional capture. These on a 3-D rendered magnetic resonance image template from the
interpretations permit to explain why parietal activ- Montreal Neurological Institute (MNI), back view. Dashed ovals
ities could also be found during pitch-related tasks cover V5 activities. The EEG electrodes of the present experiment
(Maeder et al., 2001; Rama, Paavilainen, et al., 2000; (10–20 and 10–10 electrode systems and additional intermediate
electrodes) are positioned on a realistic scalp surface (Oostenveld
Weeks et al., 1999; Zatorre, Mondor, et al., 1999; Zatorre,
& Praamstra, 2001), and head and brain contours are coregistered
Evans, et al., 1994). (rotation and scaling) to the MNI template. Electrodes corresponding
Thus, the parietal activity could be related to high- to RTPO and LTPO activities are indicated in black.
level task-related cognitive processes that would be
common to both features. In the present study, the
parietal and frontal components seem to be part of the ing the brain areas which will be later engaged in visual
network usually found to be involved in attentional mental imagery.
processes and motor preparation (Jancke, Specht, Shah,
& Hugdahl, 2003; Brunia & van Boxtel, 2001; Posner &
Temporal Components
Dehaene, 1994).
At last, sustained bilateral temporal components were
characterized by SCD polarity reversals throughout the
Temporo-Parieto-Occipital Components
stimulus duration. These activities most likely originate
By means of SCD mapping, it has also been possible to from superior temporal auditory areas (Gutschalk, Pat-
detect large and very focal bilateral TPO activities in the terson, Rupp, Uppenkamp, & Scherg, 2002) and reflect
attention condition. To our knowledge, no similar activ- auditory integration. We found these bilateral temporal
ity has ever been found in neuroimaging studies on activities for spatial and pitch processing in both tasks.
sound motion, which were generally based on passive Previous studies have suggested bilateral (Hart et al.,
listening or simple stimuli. In contrast, we used more 2004; Warren & Griffiths, 2003; Pavani et al., 2002;
complex varying stimuli in a demanding task. In the Warren et al., 2002; Zatorre, Bouffard, et al., 2002; Lewis
attention condition, to be able to follow both types of et al., 2000) or right-lateralized (Baumgart et al., 1999)
acoustic variations, most of the subjects (11 among 14) activation of the planum temporale when contrasting
asserted that they had mentally generated a visual rep- moving or spatially distributed sounds to stationary
resentation of the sound. It should be noted that TPO sounds. For pitch processing, right-lateralized (Patter-
electrode sites are close to V5 visual areas (Figure 5) son, Uppenkamp, Johnsrude, & Griffiths, 2002; Zatorre
known to be activated during visual motion perception et al., 1994) or bilateral (Hart et al., 2004; Warren &
and mental imagery (Goebel, Khorram-Sefat, Muckli, Griffiths, 2003; Thivard, Belin, Zilbovicius, Poline, &
Hacker, & Singer, 1998). The bilateral TPO activation Samson, 2000) activation in superior temporal areas
could thus correspond to visual mental imagery induced has been found when contrasting sequences with chang-
by both types of acoustic variation processing. The ing pitch to sequences with fixed pitch.
existence of these same activities with small amplitude During the varying part, left temporal activities were
before the acoustic variation onset could be associated particularly more important for pitch than for spatial
with a preparatory attentional process selectively involv- variations. This result suggests a differential processing

Bidet-Caulet and Bertrand 1697


!!     H

of spatial and pitch variations in the left temporal ‘‘where’’ acoustic features within the temporal auditory
auditory cortex. This dissociation could correspond areas.
either to an increased neuronal activity for pitch pro-
cessing or to different configurations of neural sources
for spatial and pitch processing. To our knowledge, few METHODS
studies have been interested in the difference between
pitch and spatial processing within the temporal cortex. Subjects
By contrasting sound sequences with changing pitch to Sixteen paid subjects (15 right-handers, 6 men, mean
sequences with changing location, Hart et al. (2004) and age 24 years) participated in the study and were selected
Warren and Griffiths (2003) have shown a dissociation during a selection session on the basis of their perform-
between pitch and spatial processing in the right and ance in both tasks. All were free of neurological diseases
left superior temporal planes. Nevertheless, our left- and had normal hearing. Written informed consent was
lateralized effect is consistent with Hart et al.’s additional obtained from each subject.
activity for pitch processing in the left planum polare.
According to these studies, a posteromedial (spatial)–
Task
anterolateral (pitch) dissociation would exist within the
superior temporal cortex of both hemispheres. Close Subjects had to perform two different tasks in separate
configurations of neural sources for spatial and pitch blocks. In each block, the stimuli consisted of pitch-
processing that could not be disentangled with the varying streams randomly alternating with spatially
spatial resolution of SCD computation could explain varying streams with equal probabilities. In the stream-
why we could not find any difference in the right oriented attention task (called attention task), subjects
hemisphere. were asked to follow sound variations and report its
As an interaction effect between attention and acous- final direction (high/low pitch or left/right) with a four-
tic variations has been found during the same period for direction joystick (up/down or left/right, respectively). In
the left superior component, pitch and spatial feature the diverted attention task (called control task), subjects
integration within the auditory cortex could correspond were asked to compare two successive noise-bursts
to automatic processes that are differentially modulated superimposed to the end of the stream, thus orienting
by attention. In addition, there was a small attention their attention away from the varying stream. These two
effect on the left temporal activity during the stationary bursts randomly differed in location or pitch, indepen-
part. This effect prior to stream variation could be due to dently of the underlying stream variations. Subjects had
an attention-dependent preactivation of the auditory to report with the same joystick whether the second
areas which will be subsequently involved in differential burst was to the right or to the left, or lower- or higher-
feature processing. pitched than the first one. Performances were balanced
All these modulations of the temporal activities are between tasks and also between spatial and pitch feature
consistent with a current hypothesis (Griffiths & Warren, processing with difficulty levels adapted to each subject
2002) suggesting a role of the planum temporale in differ- (see Procedure section). Each new stimulus started
entially processing auditory spectro-temporal information. between 3.3 and 4.1 sec after the subject’s response
and no specific speed instructions were given.
Conclusion
Stimuli
Combining the use of long-lasting auditory streams and
a precise spatio-temporal analysis of electrophysiologi- Auditory streams consisted of 35-msec band-passed
cal signals allows to clarify the role of the temporal, noise-bursts [5-semitone (st) wide, starting from a fre-
parietal, and frontal regions in auditory processing. The quency F, and cosine tapered] and were presented
frontal component would be involved in unspecific task- through headphones (Sennheiser PX30, Wedemark,
related processes, such as crucial stimulus expectation Germany). In both tasks, they were composed of two
and motor preparation. The parietal activity would re- successive parts, a stationary part of random duration
flect task-specific processes, such as selective attention (735, 805, 875 msec with equal probabilities) followed by
and preparation for auditory–motor transformation, an acoustically varying part (duration ranging from 980
as suggested in recent reviews (Hall, 2003; Griffiths to 2205 msec).
& Warren, 2002; Middlebrooks, 2002). The temporal Stream trajectories were piecewise-linear functions,
cortices would be more specifically involved in auditory identical for pitch and spatial variations (Figure 6A).
integration, acoustic spatial and pitch features being dif- Trajectories of pitch-varying streams were following
ferentially processed within the left temporal auditory piecewise-linear functions F(t) defined in semitones
areas. (st). These functions were equal to zero during the
The present study provides further experimental evi- stationary part and were increasing or decreasing be-
dence for a differential processing of ‘‘what’’ and tween Fm and +Fm during the varying part.

1698 Journal of Cognitive Neuroscience Volume 17, Number 11


   !           

Figure 6. Stimulus structure. (A) Stream trajectories. Auditory streams were composed of two successive parts, a stationary and a varying
part. Streams varied in location or pitch, following zigzag trajectories. Each trajectory of pitch-varying stream was following a piecewise-linear
function F(t) (gray line) defined in semitones. In the case of spatial variations, the functions A(t) were varying between Am and
+Am. These functions were identical for pitch and spatial variations. They consisted of 4 to 7 segments and could be interrupted at
variable times, either when the variations were reaching an extremum (black diamonds) or four bursts before (black circles). (B) Stream
frequency content. Gray bars represent the 35-msec, band-passed and 5-st wide noise-bursts which composed of auditory streams. In
the case of pitch-varying streams, the lower frequency F of each burst was defined in semitones by the function: F(t) = F(t) + F0 ± 1
with F(t) as described in B. The black dashed line represents the trajectory F(t)+F0, and the black dashes represent the actual lower
frequency F(t) of each burst. (C) Analysis periods. Three periods were chosen for signals analysis: stationary ([ 150, 735 msec], time 0
being the onset of the stream), varying ([ 150, 910 msec], time 0 being the onset of the varying part), and final ([ 840, 420 msec],
time 0 being the offset of the stream). They correspond to the periods having the shortest duration for each part and common to both
tasks, that is, before noise-burst (gray ovals) occurrence in the control condition. Freq = frequency; loc = location; st = semitones.

Bidet-Caulet and Bertrand 1699


!!     F

Trajectories of spatially varying streams were following both tasks, and sustained processing of the stream
piecewise-linear functions A(t) defined in degrees. trajectories was ensured in the attention task. With such
These functions were equal to 08 (ahead) during the a randomization strategy, it seems unlikely that subjects
stationary part and were increasing or decreasing be- could memorize and recognize all these trajectory pat-
tween Am and +Am during the varying part. terns during a recording session.
Depending on the difficulty level in the attention task,
Am was ranging between 308 and 608, and Fm
Procedure
between 6 and 14 st. Nevertheless, the speed of variation
remained about the same across difficulty levels (around In the first session, subjects were selected on the basis of
30 st/sec and 1608/sec). To minimize predictability, there their ability to perceive spatial variations, to perform the
were, for each difficulty level, at least six different different tasks, and to minimize blinks and eye move-
trajectories varying by the number of segments of the ments assessed by electrooculogram. First, they were
functions F(t) and A(t) (randomized between 4 and familiarized with sound sequences and tasks in a step-
7). Moreover, the trajectories were interrupted at vari- wise learning procedure using visual feedback. Then,
able times, either when the variations were reaching an they performed blocks of each task with different diffi-
extremum or four bursts before. This led to about 12 culty levels, until we could determine the difficulty level
different time courses, identical for spatial and pitch corresponding to an 80% correct response rate for each
variations, consisting of 3 to 6 direction changes for each kind of stimuli in each task (spatially or pitch-varying
difficulty level. streams in the attention task and location- or pitch-
To minimize habituation effects for both types of differing noise-bursts in the control task). Only 1 subject
variation, the lower frequency F of each burst was out of 17 had to be excluded because of excessive eye
randomly varying by ±1 st around F0, with F0 equiprob- blinks.
ably chosen between 1318, 1480, and 1661 Hz for each In the second session, subjects’ EEG were recorded
stimulus. In the case of pitch-varying streams, the lower during 12 blocks of 30 trials in both attention and
frequency of each burst was thus defined in semitones control tasks, yielding a total of 360 trials per task (180
by the function: F(t) = F(t) + F0 ± 1 with F(t) as with spatial and 180 with pitch variations). Attention (A)
described above (see Figure 6B). In the case of spatially and control (C) blocks were presented in an AA–CC–AA–
varying streams, F(t) is only varying around F0, F(t) = CC–. . . design. For each subject, the difficulty level was
F0 ± 1, throughout stimulus duration. Pitch-varying determined during the selection procedure as men-
streams were localized ahead at 08 azimuth. tioned above. In some cases, because of subjects’ per-
To give the impression of a continuously moving formance improvement or tiredness, the difficulty level
sound source, each burst was convoluted with an was adapted after each block to maintain a constant
average-listener, head-related transfer function (see performance (around 80% correct response rate)
http://sound.media.mit.edu/KEMAR.html) with a 58-step throughout the session. Subjects were instructed to
in azimuth. Compared to other studies using only in- keep their eyes open while fixating a point, and to
teraural time differences to position sounds in space refrain from blinking during stimulus presentation.
with earphones, this procedure creates a better percept
of sound objects located in the external space by mod-
EEG Recordings
ulating phase, amplitude, and frequency content of the
sounds. The subjects were seated in an armchair in a sound-
In the control condition, two additional noise-bursts attenuated and electrically shielded chamber. EEG was
(5-st wide, starting from a frequency F, and lasting recorded from 36 scalp electrodes (Easy Cap, FMS,
100 msec) appeared successively 420 and 140 msec be- Herrsching, Germany) referenced to the nose (ground
fore the end of the stream (Figure 6C). They differed electrode at forehead and impedances below 5 k)
either in location [azimuths: ±Ab; frequency: F] or in and EOG from two electrodes were placed centrally
pitch [azimuth: 08; frequency: F ± Fb]. Depending above the right eye and at its outer canthus. The elec-
on the difficulty level, Ab was ranging from 58 to 358 trode positions are shown schematically in Figure 7.
and Fb from 1 to 8 st. Because the trajectories were Signals were amplified, filtered (0.05–200 Hz band-
randomized and interrupted at variable times, subjects width), and sampled at 1000 Hz (Synamps, Neuroscan
could not predict the end of the stream, and thus, the Labs, Sterling, VA).
occurrence of the noise-bursts.
More generally, to minimize habituation and predict-
Data Analysis
ability, many acoustic parameters were randomized: the
central pitch of the stationary part (F0), the pitch of each Trials contaminated by eye blinks or excessive muscular
burst (F), and the stream trajectory. Furthermore, by activity, or corresponding to incorrect responses, were
randomizing stream duration, the subject’s attention rejected from further analysis. ERPs were averaged and
was maintained focused throughout the stimulus in digitally filtered (low-pass 35 Hz) in three time periods,

1700 Journal of Cognitive Neuroscience Volume 17, Number 11


I   !           

in the varying period, and from 840 to 420 msec in


the final period).
All signal analyses were performed with ELAN-Pack
software developed at INSERM U280.

Acknowledgments
We thank Jochen Kaiser for helpful discussions, and Pierre-
Emmanuel Aguera and Jean-François Echallier for technical
support.
Reprint requests should be sent to Aurélie Bidet-Caulet, INSERM
U280, 69675 Bron, Cedex, France, or via e-mail: bidet-caulet@
lyon.inserm.fr.

Figure 7. EEG recording sites. Electrodes were placed according


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Bidet-Caulet and Bertrand 1703


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9252 • The Journal of Neuroscience, August 29, 2007 • 27(35):9252–9261

Behavioral/Systems/Cognitive

Effects of Selective Attention on the Electrophysiological


Representation of Concurrent Sounds in the Human
Auditory Cortex
Aurélie Bidet-Caulet,1,2,3 Catherine Fischer,1,2,3,4 Julien Besle,1,2,3 Pierre-Emmanuel Aguera,1,2,3 Marie-Helene Giard,1,2,3
and Olivier Bertrand1,2,3
1 Institut National de la Santé et de la Recherche Médicale, Unite 821, Lyon F-69500, France, 2Institut Fédératif des Neurosciences, 3Université Lyon 1, and
4 Department of Functional Neurology and Epileptology, Neurological Hospital, Lyon F-69000, France

In noisy environments, we use auditory selective attention to actively ignore distracting sounds and select relevant information, as during
a cocktail party to follow one particular conversation. The present electrophysiological study aims at deciphering the spatiotemporal
organization of the effect of selective attention on the representation of concurrent sounds in the human auditory cortex. Sound onset
asynchrony was manipulated to induce the segregation of two concurrent auditory streams. Each stream consisted of amplitude modu-
lated tones at different carrier and modulation frequencies. Electrophysiological recordings were performed in epileptic patients with
pharmacologically resistant partial epilepsy, implanted with depth electrodes in the temporal cortex. Patients were presented with the
stimuli while they either performed an auditory distracting task or actively selected one of the two concurrent streams. Selective attention
was found to affect steady-state responses in the primary auditory cortex, and transient and sustained evoked responses in secondary
auditory areas. The results provide new insights on the neural mechanisms of auditory selective attention: stream selection during sound
rivalry would be facilitated not only by enhancing the neural representation of relevant sounds, but also by reducing the representation
of irrelevant information in the auditory cortex. Finally, they suggest a specialization of the left hemisphere in the attentional selection of
fine-grained acoustic information.
Key words: auditory cortex; attention; intracranial EEG; human; event-related; steady-state; concurrent sound

Introduction influence of active selection during the perception of overlapping


In ecological situations, we are often confronted with a mixture binaural streams. The main reason is the difficulty to dissociate
of sounds and it is crucial to be able to select relevant information the neural activity specifically corresponding to either sound
and resist auditory distracters for further cognitive processing stream. One way to solve this issue is to use long-duration sounds
and adapted behavioral response. The electrophysiological at different amplitude modulation frequencies. Indeed, each of
mechanisms of auditory selective attention have been extensively these sounds would elicit an evoked electrophysiological activity,
investigated through selective dichotic paradigms (for review, see named steady-state response (SSR), which has the particularity to
Giard et al., 2000). Selective attention has been shown to modu- be at the same frequency as the amplitude modulation of the
late the processing of both relevant (Hillyard et al., 1973; sound.
Woldorff and Hillyard, 1991) and irrelevant stimuli (Donald, The present electrophysiological study aims at deciphering
1987; Michie et al., 1993), with effects at multiple levels of sensory the spatiotemporal organization of the effect of selective atten-
analysis including the auditory cortex (Pugh et al., 1996; Jancke et tion on the representation of concurrent sounds in the human
al., 1999), the brainstem (Lukas, 1980, 1981) and down to the auditory cortex. Two concurrent streams at different carrier fre-
cochlea (Giard et al., 1994). quencies and amplitude modulation frequencies (21 and 29 Hz)
In a real cocktail party situation, however, the sounds do not (see Fig. 1) were used. The perceptual segregation of these two
reach the ears separately like in the dichotic paradigms classically simultaneous streams was induced by sound onset asynchrony
used in the previous studies of selective auditory attention. To (Bregman, 1990; Darwin et al., 1995; Turgeon et al., 2002): the 21
our knowledge, no electrophysiological study has investigated the Hz stream always started before the 29 Hz stream. Intracranial
electrophysiological (EEG) recordings were performed in epilep-
Received March 29, 2007; revised June 19, 2007; accepted July 11, 2007. tic patients with pharmacologically resistant epilepsy, implanted
We thank Rhodri Cusack for constructive discussion. with multicontact depth electrodes in the temporal cortex. Pa-
Correspondence should be addressed to A. Bidet-Caulet, Institut National de la Santé et de la Recherche Médicale, tients were required either to focus their attention away from the
Unite 821, Brain Dynamics and Cognition, Centre Hospitalier le Vinatier, 95 Boulevard Pinel, 69500 Bron, France.
E-mail: bidet-caulet@lyon.inserm.fr.
streams [control (C) condition] or to actively select the 21 Hz
DOI:10.1523/JNEUROSCI.1402-07.2007 stream (AS21 condition) or the 29 Hz stream (AS29 condition).
Copyright © 2007 Society for Neuroscience 0270-6474/07/279252-10$15.00/0 This paradigm therefore allowed us to compare the electro-
#         IF

Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing J. Neurosci., August 29, 2007 • 27(35):9252–9261 • 9253

(stim-part2) was equiprobably chosen between


0.810, 0.905, 1, or 1.095 s (1.095, 1.190, 1.286,
or 1.381 s for the first patient) at each trial. All
component onsets and offsets were linearly
ramped during 10 ms. Because the two streams
started at different instants (onset asynchrony),
auditory stream segregation was induced and
two distinct streams were perceived.
There were three attention conditions in sep-
arate blocks. In the first one (control condi-
tion), the patients had to detect rare noise
bursts (targets) superimposed to the stimuli
and, thus, orient their attention away from the
stimulus content. They were instructed to an-
swer as soon as they heard a noise burst, by
Figure 1. Stimuli. The stimuli were composed of two acoustic streams: a 21 Hz stream and a 29 Hz stream. The 21 Hz stream pressing a button. The superimposed target
(black bars) was composed of two amplitude-modulated tones separated by two octaves, the carrier frequency of the lower one sounds were 150 ms bandpass-filtered noise-
being equiprobably chosen between 659, 698, 740, or 784 Hz. These two tones were both amplitude modulated in phase at a bursts (20 semitones wide, starting at 784 Hz
frequency of 21 Hz. The 29 Hz stream (gray bar) consisted of one tone separated by one octave from each tone of the first stream. with 10 ms rise/fall times). The targets were de-
This tone was amplitude modulated at a frequency of 29 Hz. The 21 Hz stream always started 810 ms before the 29 Hz stream livered in 15% of the stimuli and randomly oc-
(stim-part1). After the onset of the 29 Hz stream, starts a time period of sound rivalry (stim-part2) of 810 –1095 ms duration. curred during the stimulus, 0.2, 0.5, 1.2, or 1.5 s
Because the two streams started at different times (onset asynchrony), auditory stream segregation was induced and two distinct before the end of the stimulus. When a target
streams could be perceived. In the two stream-selection attention tasks (AS21 and AS29), the stimuli included an additional part was present in a trial, the next stimulus started
(part 3) with a duration of 400 or 700 ms, during which one stream or both were changing in spatial direction. between 0.7 and 0.1 s after the patient’s re-
sponse, otherwise the intertrial interval was
randomized between 0.9 and 1.4 s.
physiological responses to acoustically identical stimuli in three In the other two conditions, the stimuli included an additional, third
different attentional contexts and to characterize the effect of part. During this last part, one stream or both were changing in spatial
selective attention before and during the sound rivalry. First, we direction. Patients were instructed to attend to a given stream (the 21 or
analyzed the transient and sustained evoked responses that are 29 Hz stream) and to indicate its final direction (left or right) with a
well known to be modulated by attention (for review, see Picton two-direction joystick. Thus, these tasks corresponded to the attentional
et al., 1978; Giard et al., 2000). Second, thanks to the distinct selection of one of the two streams (attend to the 21 Hz stream, AS21
condition; attend to the 29 Hz stream, AS29 condition). Interaural in-
amplitude modulation frequencies of the two simultaneous
tensity differences (IIDs) were used to give a realistic impression of spa-
streams, the steady-state responses were used for tagging the elec- tially moving streams during the last part of the stimulus. This part lasted
trophysiological activity corresponding to each stream indepen- 0.4 and 0.7 s for AS21 and AS29 conditions, respectively. Thus, in total,
dently. This approach thus allowed us to describe the multiple the stimuli were lasting between 1.620 and 1.905 s in the C condition,
neural mechanisms of selective attention that may operate at dis- between 2.020 and 2.305 s in the AS21 condition, and between 2.320 and
tinct processing levels and in different areas of the auditory cortex 2.605 s in the AS29 condition. In these two conditions, in 50% of the
(Heschl’s gyrus, planum temporale, and polare). trials, only the attended stream was changing in spatial direction (25% to
the left and 25% to the right), and in the other 50%, both streams were
Materials and Methods spatially changing in opposite directions (in 25%, the attended stream
Patients. We recorded the data from 12 patients (eight female and four changed to the left and the unattended to the right, and in 25% the
male, age ranging from 21 to 48 years) suffering from pharmacologically attended stream changed to the right and the unattended to the left). The
resistant partial epilepsy and candidate for surgery. Because the location four categories of stimuli were randomly presented. The level of difficulty
of the epileptic focus could not be identified using noninvasive methods, was adjusted in each patient by choosing different values of IIDs gener-
they were stereotactically implanted with multicontact depth probes. ating different ranges of spatial motion. In most of the cases, the easiest
Electrophysiological recording is part of the brain functional evaluation level (for which the motion appears to end in one ear only) was chosen to
that is performed routinely before epilepsy surgery in these patients. obtain a good rate of correct responses. The next trial started between 1.5
According to the French regulations concerning invasive investigations and 1.7 s after the patient’s response. These tasks were quite difficult.
with a direct individual benefit, patients were fully informed about the Although all the patients could perform the AS21 task, only two patients
electrode implantation, stereotactic EEG and evoked-potential record- (patients 4 and 10) could correctly perform the AS29 task.
ings, and the cortical-stimulation procedures used to localize the epilep- The intensities of all tones were corrected according to their carrier
togenic and functional brain areas. All patients gave their informed con- frequency [see Botte et al. (1989), their Fig. 1.2] and then 21 and 29 Hz
sent to participate in the experiment. The signals described here were streams were matched in intensity. Stimuli were delivered at an intensity
recorded away from the seizure focus. Several days before EEG record- level judged comfortable by the patient before the beginning of the ex-
ings, antiepileptic drugs administered to the patients had been either periment. Noise bursts were 5 dB above the stimulus intensity level.
discontinued or drastically reduced. No patient was administered with Stimuli were presented to patients in two blocks of 80 trials each for the
benzodiazepines. None of the patients reported any auditory complaint. C task and in four blocks of 40 trials for the AS21 and AS29 tasks (result-
Stimuli and task. Stimuli were composed of two acoustic streams (Fig. ing in 160 repetitions in each condition). Stimulus duration, carrier fre-
1): a 21 Hz stream and a 29 Hz stream. The 21 Hz stream was composed quency, and noise burst occurrence or final spatial direction were ran-
of two amplitude-modulated tones separated by two octaves, the carrier domized to limit habituation and predictability.
frequency of the lower one being equiprobably chosen between 659, 698, EEG recording and signal analysis. Intracranial recordings were per-
740, or 784 Hz at each trial. These two tones were both amplitude mod- formed at the Functional Neurology and Epilepsy Department (Lyon
ulated in phase at a frequency of 21 Hz. The 29 Hz stream consisted of one Neurological Hospital, Lyon, France). EEG recordings were made from
tone separated by one octave from each tone of the first stream. This tone 64 or 128 intracranial electrode contacts referenced to an intracranial
was amplitude modulated at a frequency of 29 Hz. The 21 Hz stream contact away from the superior temporal cortex. The ground electrode
always started first (stim-part1; lasting 0.810 or 1 s for the first patient). was at the forehead. Signals were amplified, filtered (0.1–200 Hz band-
Then, the 29 Hz stream started and the duration of the sound rivalry width), and sampled at 1000 Hz (Synamps; Neuroscan Labs, Sterling,
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9254 • J. Neurosci., August 29, 2007 • 27(35):9252–9261 Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing

VA) for the first six patients and were amplified, filtered (0.1–200 Hz
bandwidth), and sampled at 512 Hz (Brain Quick System Plus; Mi-
cromed, Treviso, Italy) for the next six patients.
The analysis was restricted to the electrodes located in the temporal
cortex and its immediate vicinity. Raw data were visually inspected and
trials showing epileptic spikes were discarded. Because the superior tem-
poral cortex was subsequently found to be the location of their epileptic
focus, two patients were excluded from all analysis. The evoked sustained
and transient responses from patient 3 were also excluded from analysis
because of excessive epileptic spikes, but the data were kept for the SSR
analysis because their frequency bands were less contaminated by the
spectral content of the epileptic spikes. In the AS29 condition, only the
data from two patients (4 and 10) who could perform the task were kept
for analysis. In the C condition, trials that included a target occurring
before 1.620 s after stimulus onset (⬃12% of all trials) were not analyzed.
In all conditions, trials with a motor response occurring before 1.620 s,
with a false alarm, or with an incorrect response were rejected from
additional analysis. The mean numbers of correct and nonartifacted tri-
als were 142, 136, and 123 of 160 in the C, AS21, and AS29 conditions,
respectively. The EEG of four patients (6, 7, 8, and 9) were notch-filtered
at 50 Hz because of excessive power line artifact.
Evoked responses were averaged over a 2 s period and corrected with
respect to the same baseline defined between ⫺150 and 0 ms before
stim-part1 onset.
The periodic SSR, evoked by the amplitude modulation of the streams
at 21 or 29 Hz was analyzed by means of a wavelet decomposition, which
provides a good compromise between time and frequency resolutions.
Each single-trial signal was transformed in the time-frequency (TF) do- Figure 2. Delineation of the main auditory cortical regions on a top view of the superior
main by convolution with complex Gaussian Morlet’s wavelets with a temporal plane after 3D rendering of the cortical surface of the MNI standard brain. This sche-
ratio f/␴f of 10, with f being the central frequency of the wavelet and ␴f its matic representation will be used in Figures 4 – 8.
SD (for details, see Tallon-Baudry and Bertrand, 1999). A baseline cor-
rection was applied on TF plots by subtracting, in each frequency band, For each EEG component, a first statistical analysis in each of the 10
the prestimulus power between ⫺250 and ⫺150 ms before stimulus patients was performed by directly comparing the C and AS21 conditions
onset. From this TF analysis, time profiles of the 21 Hz and 29 Hz SSR using Mann–Whitney tests. In patients 4 and 10, we further compared
could be computed and were used for statistical analysis. the three conditions using a global Kruskal–Wallis test followed by two-
Statistical analysis. Statistical analysis in the three conditions focused by-two Mann–Whitney post hoc tests.
on several electrophysiological components: the transient responses Only those effects are discussed that met Bonferroni-corrected p value
evoked by the onsets of stim-part1 and stim-part2, the sustained re- criteria. In each patient, the probability threshold of 0.05 was thus di-
sponses elicited during stim-part1, and the SSR at 21 and 29 Hz during vided by the number of tests performed (i.e., the number of time win-
both parts of the stimulus (restricted to the shortest duration of stim- dows tested over all electrode contacts investigated).
part1 and stim-part2, i.e., 0.810 s for each part). Because the data were All signal analyses were performed using the ELAN-Pack software
not normally distributed, only nonparametric tests (Wilcoxon or Mann– developed at Institut National de la Santé et de la Recherche Médicale,
Whitney) were used. Unite 821.
Statistical analysis of the sustained responses was performed on the Electrode implantation, anatomical registration, and normalization.
0.2–15 Hz bandpass-filtered EEG to remove the SSR. Because the tran- Electrode contacts were 2 mm long and spaced every 3.5 mm (center to
sient responses to stim-part2 onset could be direct-current shifted by the center). Depth probes (diameter, 0.8 mm) with 10 or 15 contacts each
ongoing sustained response (of stim-part1), statistical analysis of the were inserted perpendicularly to the sagittal plane using Talairach’s ste-
transient evoked responses was performed on the 2–150 Hz bandpass- reotactic grid (Talairach and Tournoux, 1988). Numbering of contacts is
filtered EEG. To identify in each patient the electrode contacts where a increasing from medial to lateral along an electrode track. Electrode
transient or sustained response was emerging, a time-varying Wilcoxon locations were measured on x-ray images obtained in the stereotactic
test was computed from the single trials. It was applied to the mean frame. The depth of penetration of each contact was measured on the
amplitude of successive 20 ms time windows between 0 and 300 ms frontal x-ray image from the tip of the electrode to the midline, which
(transient responses) and successive 100 ms windows between 300 and was visualized angiographically by the sagittal sinus. The coregistration
800 ms (sustained responses) after the onset of each stimulus part, com- of the lateral x-ray image and a midsagittal magnetic resonance imaging
pared with a prestimulus baseline (defined between ⫺100 and 0 ms (MRI) scan, both having the same scale of 1, allowed us to measure the
before stim-part1 onset). For contacts that showed a significant emerging electrode coordinates in the individual Talairach’s space defined by the
response, differences between conditions were estimated by Mann– median sagittal plane, the anterior commissure–posterior commissure
Whitney or Kruskal–Wallis tests applied to the same windows as de- (AC–PC) horizontal plane, and the vertical AC frontal plane, these ana-
scribed for the response identification test. tomical landmarks being identified on the three-dimensional (3D) MRI
The emergence of SSR was assessed by Wilcoxon tests comparing the scans. With this procedure, we could superpose each electrode contact
mean power of each frequency (21 and 29 Hz) over successive 100 ms onto the patients’ structural MRI scans. The accuracy of the registration
windows between 0 and 800 ms after the onset of each stimulus part to procedure was 2 mm, as estimated on another patient’s MR images ob-
the prestimulus baseline power (between ⫺250 and ⫺150 ms before tained just after electrode explantation and in which electrode tracks
stim-part1 onset) of the respective frequency (note that the prestimulus were still visible.
baseline is shifted away from 0 ms because wavelet analysis tends to Four patients were implanted in the right hemisphere only, two in the
stretch out the early poststimulus low frequency components). For con- left only and four in both hemispheres. In all implanted hemispheres, at
tacts that showed a significant SSR, differences between conditions were least one electrode track was located in the superior temporal cortex.
estimated by Mann–Whitney or Kruskal–Wallis tests applied to the same Electrodes H and H⬘ (prime denoting the left hemisphere) were posi-
windows as described for the response-identification test. tioned posteriorly, passing through Heschl’s gyrus (HG), the planum
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Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing J. Neurosci., August 29, 2007 • 27(35):9252–9261 • 9255

patients could perform the AS21 task with


at least 85% of correct responses (mean,
97.2%), only two patients could reach the
same percentage of correct responses in
the AS29 task (patient 4, 85%; patient 10,
88.1% of correct responses).
The difference of performance between
the two stream-selection tasks could be at-
tributable to the asymmetric construction
of the stimuli. Indeed, the 21 Hz stream
could be easier to select because it con-
tained the lowest and highest frequencies
of the signal and not the middle one, be-
cause it started first, and because it con-
tained two frequencies and not only one.
This difference of performance proba-
bly reflects a greater attentional load in the
AS29 than in the AS21 condition. This dif-
ference is likely to favor the detection of
attentional effects on the electrophysiolog-
ical responses in the AS29 condition.
In the following, for each EEG compo-
nent, the results of statistical analysis com-
paring the C and AS21 conditions in 10
(for steady-sate responses) or 9 patients
(for transient and sustained responses, re-
spectively) will be presented first, followed
by the results of a second analysis consid-
Figure 3. Illustration of the typical electrophysiological responses and their location in the 3D rendering of the temporal cortex ering the three conditions in patients 4 and
of patient 10. A, B, The evoked response analysis consists of averaging single trials from EEG filtered with different bandwidth to 10.
dissociate transient and sustained responses. Top curves, Unfiltered evoked responses; middle curves, sustained responses ob-
tained from 0.2–15 Hz filtered EEG; bottom curves, transient responses obtained from 2 to 150 Hz filtered EEG. The periodic
Electrophysiological responses
steady-state activity is visible on the unfiltered and transient responses in A. C, The SSR analysis is based on a time-frequency
Distinct electrophysiological components
transformation of electrophysiological activities (time-frequency power averaged after a wavelet-based transform of each single
were identified in different sites of the au-
trial). Twenty-one, 29, and even 42 Hz steady-state evoked activities are clearly visible on the time-frequency plots. The time
ditory cortex. Figure 3 illustrates these
profiles of SSR power could then be constructed at 21 and 29 Hz from the time-frequency plots. All of these responses are baseline
corrected with respect to a prestimulus period preceding the stimulus onset. components on the 3D rendering of pa-
tient 10’s right temporal cortex. The
evoked responses are shown after single
temporale (PT), and the superior temporal gyrus (STG), and electrodes trial averaging. The transient and sus-
T, T⬘, and W were positioned anteriorly, passing through the HG and the tained responses (Fig. 3 A, B) could be dissociated by digitally
anterior PT or the planum polare (PP). Electrodes A, A⬘, B, B⬘, and C filtering the EEG signal using two different bandwidth, 2–150 and
were penetrating through the middle temporal gyrus (MTG). Electrodes
0.2–15 Hz, respectively. The periodic SSRs clearly visible on the
N and N⬘ were located just above the superior temporal plane, in the
parietal operculum. Although intracranial recordings in epileptic pa- (unfiltered or transient) evoked responses were further analyzed
tients provide a sparse spatial sampling of the auditory cortex, we could using a time-frequency decomposition of the electrophysiologi-
access not only primary auditory areas (posteromedial part of the HG), cal signals (time-frequency power averaged after a wavelet-based
but also posterior and anterior secondary auditory regions (PT and PP). decomposition of each single trial). The time profiles of the SSR
The electrode coordinates of each patient were converted from the power could then be constructed at 21 and 29 Hz from the time-
individual Talairach’s space to the normalized Talairach’s space (Ta- frequency plots (Fig. 3C).
lairach and Tournoux, 1988), and then to the Talairach’s space of the
All of these components were present during both stim-part1
Montreal Neurological Institute (MNI) standard brain. Eventually, elec-
trode contacts and experimental effects of all patients were plotted on a and stim-part2. Effects of attention on these electrophysiological
3D rendering of the temporal cortices of the MNI standard brain (corti- components could thus be investigated during both stimulus
cal surface segmentation by FreeSurfer software, http://surfer.nmr.mgh- parts, because the inputs were acoustically identical and only the
.harvard.edu). This procedure facilitated the comparison across patients focus of attention was changing. These attentional effects will be
of the activated sites that could be positioned with respect to the main presented, for the group of patients, on a schematic view of the
superior temporal structures that were delineated on the standard brain superior temporal cortex constructed from the MNI standard
(Fig. 2).
brain (Fig. 2). For sake of clarity, we only reported, in Figures
Results 4– 8, electrodes passing through the superior temporal cortex and
Behavioral data the attention effects found at these electrodes. These plots show
The control task presented no major difficulty, but was demand- that the auditory cortex could be covered with a good spatial
ing enough to keep the patients alert (mean, 99.3% of correct sampling over the group of patients. To illustrate the time course
responses and 0.2% of false alarms). The stream-selection atten- of the responses, typical waveforms have been plotted from elec-
tion tasks (AS21 and AS29) were more difficult. Although all the trode contacts showing attentional effects.
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9256 • J. Neurosci., August 29, 2007 • 27(35):9252–9261 Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing

Transient evoked responses


(nine patients)
We found significant transient evoked re-
sponses after both stim-part1 and stim-
part2 onsets at most contacts of electrodes
H, H⬘, T, T⬘, and W (199 contacts of 208
tested contacts). Three main transient
waves could be observed: a first one maxi-
mal between 30 and 75 ms, a second one
⬃100 ms (between 75 and 150 ms), and a
third one starting ⬃150 –170 ms. These
waves had smaller amplitudes and slightly
later latencies after stim-part2 onset com-
pared with stim-part1 onset.
After stim-part1 onset (onset of the 21
Hz stream), transient waves were found
modulated by attention at 67 contacts (of
199) across the nine patients kept for this
analysis (Fig. 4). These waves had signifi-
cantly larger amplitudes in the AS21 than
in the C condition at all the 67 contacts.
The three waves could be affected by atten-
tion, but most effects were found on the Figure 4. Attention effects on the transient evoked responses during stim-part1. Three transient evoked waves were found
second wave (between 75 and 150 ms) modulated by attention after stim-part1 onset: a first one between 30 and 75 ms (yellow), a second one between 75 and 150 ms
(green) and a third one between 150 and 280 ms (blue). Ovals on the anatomical plots correspond to contacts where at least the
(Fig. 4, green ovals) in several sites of the
AS21 ⬎ C effect was found, whereas diamonds correspond to contacts where any other effect (not the AS21 ⬎ C effect) was
auditory cortex (HG, PP, and PT) in both found. The time courses of these effects are depicted on the left and right sides of the figure. Evoked potentials waveforms were
hemispheres. For the two patients who obtained by averaging the single trials from the 2–150 Hz filtered EEG. Transient evoked responses with significant difference
could perform the AS29 task, the transient between conditions are indicated by asterisks. The black, gray, and hatched boxes indicate which conditions are compared.
waves were found modulated by attention
at 10 contacts (of 36). The main effects
were larger amplitudes in the AS21 than in
the C condition at nine contacts, in the
AS21 than in the AS29 condition at six
contacts, and in the AS29 than in the C
condition at eight contacts. In other
words, we globally found larger ampli-
tudes in the AS21 than in the AS29, and, in
turn, greater than in the C condition (in
the rest of the study, this will be formu-
lated as AS21 ⬎ AS29 ⬎ C).
To summarize, the main effects of at-
tention on the transient evoked responses
to the onset of the 21 Hz stream (AS21 ⬎
AS29 ⬎ C) strongly suggest the existence
of two levels of attentional selection: a first
one to select amplitude modulated
streams from target sounds (AS21 ⬎ C and
AS29 ⬎ C) and a second to select one of Figure 5. Attention effects on the transient evoked responses during stim-part2. Three transient evoked responses were found
modulated by attention after stim-part2 onset: a first one between 30 and 75 ms (yellow), a second one between 75 and 150 ms
the two streams (AS21 ⬎ AS29). These ef- (green), and a third one between 150 and 280 ms (blue). Squares on the anatomical plots correspond to contacts where at least the
fects show that selective attention to audi- C ⬎ AS21 effect was found. Ovals correspond to contacts where only the AS21 ⬎ C effect was found. Diamonds correspond to
tory streams (compared with the control contacts where any other effect (not the AS21 ⬎ C or C ⬎ AS21 effects) was found. The time courses of these effects are depicted
condition) operates by enhancing the neu- on the left and right sides of the figure. The curves are obtained by averaging single trials from 2 to 150 Hz filtered EEG. Transient
ral representation of the relevant informa- evoked responses with significant difference between conditions are indicated by asterisks. The black, gray, and hatched boxes
tion: the more relevant the sound, the indicate which conditions are compared.
more increased its neural representation.
After stim-part2 onset (onset of the 29 Hz stream), the tran- and mainly in the left hemisphere. We found attentional effects in
sient waves were found modulated by attention at 34 contacts (of only one of the two patients who could perform the AS29 task, at
199) across eight of the nine patients kept for this analysis (Fig. 5). seven contacts (of 13) in the right hemisphere. For this patient,
These waves had significantly smaller amplitudes in the AS21 the main effects were larger amplitudes of the transient waves in
than in the C condition at 31 of the 34 contacts. The three tran- the C than in the AS21 condition at four contacts, in the AS21
sient waves could be affected by attention, but most effects were than in the AS29 condition at three contacts, and in the C than in
found on the third wave (between 150 and 280 ms) (Fig. 5, blue the AS29 condition at five contacts. These effects could be sum-
squares) in several sites of the auditory cortex (HG, PP, and PT), marized by C ⬎ AS21 ⬎ AS29. The transient evoked responses at
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Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing J. Neurosci., August 29, 2007 • 27(35):9252–9261 • 9257

ms) than the attentional enhancement ob-


served during stim-part1 (between 75 and
150 ms).
These transient waves were also found
at some contacts of electrodes A, A⬘, B, B⬘,
and C in the MTG, or N and N⬘ in the
parietal operculum. These responses were
present at several successive contacts with
smaller amplitudes than at electrode con-
tacts in the superior temporal cortex.
Thus, the transient waves observed in the
MTG and the parietal operculum probably
mainly reflect the diffusion by volume
conduction of generators located in the su-
perior temporal cortex. The few attention
effects found in these regions corre-
sponded closely to the effects found on the
nearest electrode in the superior temporal
Figure 6. Emergence and attention effects of the evoked sustained responses during stim-part1. The distribution of the cortex.
sustained responses is indicated by disks (red, positive potentials; green, negative potentials) on the anatomical plots. The
diameter of these disks correspond to the normalized (across all contacts of electrodes passing through the superior temporal
Sustained evoked responses
cortex in all patients) mean amplitude of the sustained waves between 450 and 750 ms. Black squares on the schema correspond
to contacts where only the C ⬎ AS21 effect was found. Black ovals correspond to contacts where only the AS21 ⬎ C effect was (nine patients)
found and gray ovals correspond to contacts where at least the AS21 ⬎ C effect was found. Diamonds correspond to contacts At most of the electrode contacts present-
where any other effect (not the AS21 ⬎ C or C ⬎ AS21 effects) was found. The time courses of these effects are depicted on the ing transient responses (179 of 199), a sus-
left and right sides of the figure (significant differences are indicated by gray shaded areas). The curves are obtained by averaging tained response was present during both
the single trials from 0.2–15 Hz filtered EEG and are baseline corrected with respect to the [⫺100, 0 ms] period preceding parts of the stimulus (Fig. 6). If the sus-
stim-part1. The black, gray, and hatched boxes indicate which conditions are compared. tained response during stim-part1 most
likely corresponds to the processing of the
21 Hz stream only, it is not possible to dis-
sociate the sustained response corre-
sponding to the processing of the 21 and
29 Hz streams, respectively, during sound
rivalry (stim-part2). Therefore, the atten-
tional effects on the sustained responses
were investigated during stim-part1 only.
The sustained responses were found
modulated by attention at 60 contacts (of
179) across six of the nine patients kept for
this analysis (Fig. 6). These responses had
significantly larger amplitudes in the AS21
than in the C condition, at 58 of the 60 con-
tacts. These effects were mainly concentrated
between 400 and 600 ms, in several areas of
the auditory cortex (HG, PP, and PT) in
both hemispheres. For the two patients who
could perform the AS29 task, the sustained
Figure 7. Emergence and attention effects of the evoked 21 Hz SSR during parts 1 and 2. The distribution of the 21 Hz SSR is waves were found modulated by attention at
indicated by gray disks on the anatomical plots. The diameter of these disks correspond to the normalized (across contacts of each 20 contacts (of 33). The main effects were
electrode passing through the superior temporal cortex in each patient) mean amplitude of the SSR between 450 and 750 ms larger amplitudes in the AS21 than in the C
(stim-part1). Black squares on the schema correspond to contacts where only the C ⬎ AS21 effect was found and gray squares
condition at 18 contacts, in the AS21 than in
correspond to contacts where at least the C ⬎ AS21 effect was found. Black ovals correspond to contacts where only the AS21 ⬎
C effect was found. Diamonds correspond to contacts where any other effect (not the AS21 ⬎ C or C ⬎ AS21 effects) was found.
the AS29 condition at six contacts, and in the
The time courses of these effects are depicted on the left and right sides of the figure (significant differences are indicated by gray AS29 than in the C condition at three con-
shaded areas). The curves are obtained by computing the time profile at 21 Hz from the wavelet-based time-frequency decom- tacts. These effects could be summarized by
position. The black, gray, and hatched boxes indicate which conditions are compared. AS21 ⬎ AS29 ⬎ C.
Thus, the attentional modulations ob-
served in the sustained responses evoked
by the 21 Hz stream (AS21 ⬎ AS29 ⬎ C)
the stim-part2 onset most likely correspond to an electrophysio- are similar to the effects found on the transient evoked responses
logical response to the onset of the 29 Hz stream. The main effect after stim-part1 onset: the more relevant the sound, the more
in the left hemisphere, AS21 ⬍ C, indicates that selective atten- increased its neural representation.
tion operates by reducing the neural representation of irrelevant As for the transient waves, the sustained responses found at
information (the 29 Hz stream in the AS21 condition). This re- some electrode contacts in the MTG and in the parietal opercu-
duction process occurred later in latency (between 150 and 250 lum mainly reflected the diffusion by volume conduction of gen-
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9258 • J. Neurosci., August 29, 2007 • 27(35):9252–9261 Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing

erators located in the superior temporal


cortex. Similarly, the effects of attention
found in these regions corresponded to the
effects found on the nearest electrode in
the superior temporal cortex.

Steady-state evoked activities


(10 patients)
SSRs were emerging at 21 and 29 Hz at
several contacts (80 and 39 of 208 contacts,
respectively) along the HG in all patients
(Figs. 7, 8, disks). They were very focal be-
cause they could not be observed either on
other electrode contacts in the superior Figure 8. Emergence and attention effects of the evoked 29 Hz SSR during stim-part2. The distribution of the 29 Hz SSR is
temporal cortex (PT or PP), or in the MTG indicated by gray disks. The diameters of these disks correspond to the normalized (across contacts of each electrode passing
or in the parietal operculum. The distribu- through the superior temporal cortex in each patient) mean amplitude of the SSR between 1100 and 1500 ms (stim-part2). Black
tions of the 21 Hz SSR were quite similar squares on the schema correspond to contacts where only the C ⬎ AS21 effect was found whereas black ovals correspond to
during stim-part1 and stim-part2. Two bi- contacts where only the AS21 ⬎ C effect was found. Diamonds correspond to contacts where at least the AS29 ⬎ C effect (but not
lateral clusters could be observed in the the AS21 ⬎ C or C ⬎ AS21 effects) was found. The time courses of these effects are depicted on the left and right sides of the figure
HG: a first one in its posteromedial part (significant differences are indicated by gray shaded areas). The curves are obtained by computing the time profile at 29 Hz from
the wavelet-based time-frequency decomposition. The black, gray, and hatched boxes indicate which conditions are compared.
and a second one in a more anterior part
(Fig. 7). The 21 and 29 Hz SSRs presented
similar distributions (Fig. 7, 8), except that
the 29 Hz SSR was absent in the anterior
HG cluster in the left hemisphere (Fig. 8).
In most of the cases, the maximum values
of the 21 and 29 Hz SSRs were at different,
but adjacent, contacts. These activities
lasted during several hundreds of millisec-
onds during each stimulus part.
During stim-part1, 21 Hz SSRs were
modulated by attention at 20 contacts (of
80) in eight patients (Fig. 7). The power of
the SSR was significantly larger in the AS21
than in the C condition at five contacts in
the left hemisphere, and significantly more
prominent in the C than in the AS21 con-
dition at 15 contacts in the right hemi-
sphere. For the two patients who could
perform the AS29 task, the 21 Hz SSRs
were found modulated by attention at six
contacts (of 14). The main effects were
AS21 ⬎ AS29 at one contact in the left
hemisphere and C ⬎ AS29 ⬎ AS21 at five
contacts in the right hemisphere. In both
hemispheres, these effects were mainly Figure 9. Schematic representation of the main attentional effects. For each electrophysiological response, the main effects
concentrated between 250 and 550 ms. between conditions and their latencies, are arbitrarily schematized with black, gray, and hatched boxes. The amount of activity is
In the left hemisphere, the main effects depicted with the same height in the control condition (C), whereas it can be enhanced or reduced in the stream-selection
on the 21 Hz SSRs (AS21 ⬎ C and AS21 ⬎ attention conditions (AS21 and AS29). Effects corresponding to an attentional facilitation are indicated by up-pointing arrows (the
representation of the relevant sound is increased). Effects corresponding to an attentional reduction are indicated by down-
AS29) again indicate that selective atten-
pointing arrows (the representation of the irrelevant sound is reduced). It is important to note that the results indicated for the
tion can operate by enhancing the neural AS29 condition correspond to two patients only, whereas results in the C and AS29 conditions correspond to 9 or 10 patients.
representation of relevant information in Before the actual sound rivalry, transient and sustained evoked responses in both hemispheres, and 21 Hz SSR in the left hemi-
the auditory cortex. In the right hemi- sphere only, are enhanced by selective attention (compared with the control condition). During the situation of sound rivalry, in
sphere, the main effect (C ⬎ AS29 ⬎ the left hemisphere, 21 Hz SSR is reduced when attention was directed to the other stream (AS29 condition). In the left hemi-
AS21) is opposed to effects observed in the sphere, transient responses are reduced when attention was directed to the other stream (AS21 condition). In both hemispheres,
left hemisphere. This rather unexpected the 29 Hz SSR is enhanced (reduced) when the 29 Hz stream is attended (ignored). During both parts, in the right hemisphere, the
result suggests a different specialization of attentional effects on 21 Hz SSR (hatched circles) are different from all other effects. This rather unexpected result suggests a
the right and left auditory cortices in atten- specialization of the right and left auditory cortices in attentional processes.
tional processes.
During stim-part2, 21 Hz SSRs were also found modulated by right hemisphere. For the two patients who could perform the
attention at six contacts (of 80) in two patients. The power of the AS29 task, the main effects were AS21 ⬎ AS29 and C ⬎ AS29 at
SSR was significantly larger in the C than in the AS21 conditions one contact in the left hemisphere and C ⬎ AS29 ⬎ AS21 at five
at one contact in the left hemisphere and at five contacts in the contacts in the right hemisphere. The effects were mainly concen-
#         

Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing J. Neurosci., August 29, 2007 • 27(35):9252–9261 • 9259

trated between 350 and 600 ms in the left hemisphere, and be- by Linden et al. (1987), attention was directed on sound fre-
tween 50 and 800 ms in the right hemisphere. quency or intensity.
Thus, in the left hemisphere, the effects on the 21 Hz SSRs Thus, it seems that attention can affect SSR in specific situa-
(AS21 ⬎ AS29 and C ⬎ AS29) show that selective attention can tions, when the task-relevant feature is the amplitude modulation
function by reducing the neural representation of the irrelevant of the sound (i.e., the very specific acoustic feature responsible for
information (the 21 Hz stream in the AS29 condition). This re- the SSR), or in more complex situations when the attentional
duction process occurred later in latency (⬃350 ms after stim- selection is difficult.
part2 onset) than the attentional enhancement observed during
stim-part1 (⬃250 ms after stim-part1 onset). The attentional Localization of auditory selective-attention effects
effect on the 21 Hz SSR in the right hemisphere (C ⬎ AS29 ⬎ In the present study, SSRs generated in the PAC, as well as SSRs
AS21) indicates that the hemispheric specialization noted above and sustained and transient waves generated in anterior or pos-
would remain during the sound rivalry. terior secondary auditory areas (PP and PT, respectively) were
All the effects on 21 Hz SSR were mainly located in the pos- modulated by selective attention. Although effects of selective
teromedial cluster of the HG. attention in nonprimary auditory areas have been observed in
During stim-part2, 29 Hz SSRs were found modulated by at- several previous experiments using positron emission tomogra-
tention at three contacts (of 39) in two patients (Fig. 8). The phy (PET) (Alho et al., 1999), functional MRI (Pugh et al., 1996;
power of the SSR was significantly larger in the AS21 than in the Jancke et al., 1999), MEG (Woldorff et al., 1993; Ahveninen et al.,
C condition at one contact in the left hemisphere, and in the C 2006), or cortical EEG (Neelon et al., 2006), only few neuroim-
than in the AS21 condition at one contact in each hemisphere. In aging studies have reported attention effects in the primary audi-
the two patients who could perform the AS29 task, the main tory cortex (Fujiwara et al., 1998; Alho et al., 1999; Jancke et al.,
effects were AS29 ⬎ C and AS29 ⬎ AS21 at three contacts in each 1999). The present results confirm that selective auditory atten-
hemisphere (of seven). The effects were mainly concentrated be- tion can alter the sensory responses in both the primary and
tween 300 and 500 ms in both hemispheres. associative auditory areas. These attentional modulations in sen-
Thus, the main condition effects, in both hemispheres, on the sory areas could be controlled by the frontal cortex. Indeed, pre-
29 Hz SSRs (AS29 ⬎ C ⬎ AS21) would confirm that selective frontal lesion has been shown to increase early auditory evoked
attention can both enhance the neural representation of relevant potentials to irrelevant sounds (Knight et al., 1989) and frontal
sounds (the 29 Hz stream in the AS29 condition) and reduce the regions would be activated according to the amount of attention
neural representation of irrelevant information (the 29 Hz required to perform an auditory task (Jancke and Shah, 2002).
stream in the AS21 condition).
Neural mechanisms of auditory selective attention
The present experiment manipulated two levels of selection: a
Discussion first level to select targets from amplitude-modulated streams
The present results provide both a precise time course and a (control condition), and a second level to select one of the two
detailed localization of the effects of selective attention on the auditory streams (AS21 and AS29 conditions). Therefore, in the
representation of concurrent sounds in the human auditory cor- control condition, the same amount of attention was devoted to
tex. They demonstrate that steady-state responses, generated in both streams, whereas in the other two attentional conditions,
the primary auditory cortex, can be modulated by selective atten- one auditory stream was attended and the other ignored. The
tion whereas the transient and sustained evoked responses are main effects of selective attention are summarized in Figure 9 and
rather modulated in anterior and posterior associative auditory interpreted below. It can be noticed that effects on the 21 Hz SSRs
areas. They also provide new insights on the neural mechanisms in the right hemisphere present a particular pattern (hatched
of selective auditory attention. circles) that will be discussed in terms of hemispheric
specializations.
Attentional modulation of SSRs
Twenty-one and 29 Hz SSRs were found bilaterally along the HG Attentional effects on relevant and irrelevant
in two foci: in the posteromedial part of the HG, corresponding sound representation
to the primary auditory cortex (PAC) (Liegeois-Chauvel et al., Before the period of sound rivalry (Fig. 9A), selective attention
1991; Rivier and Clarke, 1997), and in a more anterior part of the seems to operate only by enhancing the neural representation
HG, considered an associative auditory area. An origin of the (transient, sustained, and 21 Hz SSR responses) of the relevant
steady-state activities in two foci of the HG is consistent with sound (21 Hz stream), as early as 75 ms after stimulus onset. This
previous magnetoencephalographic (MEG) (Gutschalk et al., mechanism of attentional facilitation has been observed in pre-
1999) and intracranial EEG (Liegeois-Chauvel et al., 2004) vious electrophysiological studies on transient waves (Hillyard et
studies. al., 1973; Woldorff and Hillyard, 1991; Alcaini et al., 1995), sus-
Very few studies have investigated the effect of attention on tained responses (Picton et al., 1978), and SSRs (Ross et al., 2004).
SSR and this issue is still debated. We found that SSR can be During sound rivalry (Fig. 9B), selective attention seems to
altered according to the orientation of attention in both foci of mainly operate by reducing the neural encoding of the irrelevant
the HG. Our results differ from some previous observations (Lin- sounds (21 Hz SSR in the AS29 condition; and transient re-
den et al., 1987), but are consistent with others (Ross et al., 2004). sponses to 29 Hz stream onset and 29 Hz SSR in the AS21 condi-
This discrepancy may be explained by the different tasks used in tion) from 150 ms after stimulus onset. This reduction of irrele-
these studies. In the present experiment, attention is manipulated vant information processing is consistent with previous
during a situation of sound rivalry, which requires strong atten- electrophysiological (Alho et al., 1987, 1994; Donald, 1987;
tional effort to select the relevant amplitude modulated stream. Michie et al., 1990, 1993; Alain and Woods, 1994), PET (Ghatan
In the study by Ross et al. (2004), attention was directed on the et al., 1998; Kawashima et al., 1999) and lesion (Knight et al.,
amplitude modulation of the sounds whereas, in the experiment 1989) studies on auditory selective attention. As shown previ-
   #            

9260 • J. Neurosci., August 29, 2007 • 27(35):9252–9261 Bidet-Caulet et al. • Selective Attention in Concurrent Sound Processing

ously (Donald, 1987; Michie et al., 1990, 1993), this reduction neural activity in the left auditory cortex. In the control condi-
occurred later in the sensory processing chain than the atten- tion, a more global analysis of the sounds was sufficient to detect
tional enhancement. Thus, attentional facilitation and inhibition superimposed noise bursts and, thus, preferentially enhanced the
are most likely to be distinct processes with different temporal neural activity in the right auditory cortex.
properties. In addition, the inhibition mechanisms seem to pref-
erentially occur in situation of sound rivalry. This agrees with Conclusion
Lavie’s (2005) proposition that only high perceptual load can The present results demonstrate that selective auditory attention
prevent distracter processing. can (1) affect the evoked responses in the primary and associative
To summarize, our findings show that in case of difficult at- auditory areas, (2) modulate the steady state responses in a situ-
tentional selection, mechanisms of active rejection of irrelevant ation of sound rivalry, (3) operate by enhancing the neural rep-
sounds complement those of enhancement of relevant informa- resentation of relevant sounds, and (4) also function by reducing
tion processing, probably to improve the neural signal-to-noise the neural representation of irrelevant information when sound
ratio of the attended input. competition makes the attentional selection difficult. Finally, our
data suggest a specialization of the left auditory cortex in the
Sensory filtering model of auditory attention attentional selection of fine-grained acoustic information.
The transient (negative or positive) waves elicited after stream
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