Auditory Peripersonal Space in Humans
Alessandro Farnè1,2 and Elisabetta Làdavas1,3
Abstract
& In the present study we report neuropsychological evi-
dence of the existence of an auditory peripersonal space
representation around the head in humans and its character-
istics. In a group of right brain-damaged patients with tactile
extinction, we found that a sound delivered near the
ipsilesional side of the head (20 cm) strongly extinguished a
tactile stimulus delivered to the contralesional side of the head
(cross-modal auditory – tactile extinction). By contrast, when
an auditory stimulus was presented far from the head (70 cm),
cross-modal extinction was dramatically reduced. This spatially
specific cross-modal extinction was most consistently found
(i.e., both in the front and back spaces) when a complex sound
was presented, like a white noise burst. Pure tones produced
spatially specific cross-modal extinction when presented in the
back space, but not in the front space. In addition, the most
severe cross-modal extinction emerged when sounds came
INTRODUCTION
Similar to other psychological domains, research in
neuropsychology has historically focused on a single
sensory modality at a time. However, we typically receive
a simultaneous flow of information from each of our
different senses in real word situations. Therefore, our
perception of objects in the world is the product of an
integrated multisensory processing.
Some of the sensory integrated systems responsible of
such processing have now been documented physiolog-
ically, and their relevance for the coding of the space
has been shown. For instance, animal experiments have
revealed brain areas specialized for the coding of visual
space surrounding the body (peripersonal space).
Neurons have been reported in parietal and frontal
lobes that effectively respond only when visual stimuli
are located in spatial proximity to a particular body part
(e.g., face and hand). In particular, these bimodal
neurons located in different brain structures, such as
the putamen (Graziano & Gross, 1993), the ventral
premotor cortex (areas F4) (Rizzolatti, Scandolara,
Matelli, & Gentilucci, 1981), the parietal areas 7b
1
CNC Centro Neuroscienze Cognitive, Dipartimento di Psico-
logia, Università di Bologna, 2S. Camillo Hospital, Venice, Italy,
3
INRCA Hospital ‘‘I Fraticini,’’ Florence, Italy
D 2002 Massachusetts Institute of Technology
from behind the head, thus showing that the back space is
more sensitive than the front space to the sensory interaction
of auditory – tactile inputs. Finally, when cross-modal effects
were investigated by reversing the spatial arrangement of
cross-modal stimuli (i.e., touch on the right and sound on the
left), we found that an ipsilesional tactile stimulus, although
inducing a small amount of cross-modal tactile – auditory
extinction, did not produce any spatial-specific effect. There-
fore, the selective aspects of cross-modal interaction found
near the head cannot be explained by a competition between a
damaged left spatial representation and an intact right spatial
representation. Thus, consistent with neurophysiological
evidence from monkeys, our findings strongly support the
existence, in humans, of an integrated cross-modal system
coding auditory and tactile stimuli near the body, that is, in the
peripersonal space. &
(Graziano & Gross, 1995), and ventral intraparietal areas
(VIP; Duhamel, Colby, & Goldberg, 1991), have tactile
receptive fields (RFs) on the hand or on the face and
correspondent visual RFs in the space immediately
adjacent the tactile fields. These neurons typically
respond best, that is, with the highest firing rate, to
visual stimuli presented near the skin surface. In con-
trast, weak responses are evoked when the same visual
stimuli are presented far from the skin (Duhamel, Colby,
& Goldberg, 1998; Graziano, Hu, & Gross, 1997; Fogassi
et al., 1996; Gentilucci, Scandolara, Pigarev, & Rizzolatti,
1983; Gentilucci et al., 1988).
In close analogy with monkey data, recent neuro-
psychological findings from our laboratory provided
the first evidence that the human brain forms integrated
visual – tactile representations of the peripersonal space
surrounding the hand (Làdavas, Farnè, Zeloni, & di
Pellegrino, 2000; Làdavas, di Pellegrino, Farnè, & Zeloni,
1998; di Pellegrino, Làdavas, & Farnè, 1997) and the face
(Làdavas, Zeloni, & Farnè, 1998). In these studies, a
cross-modal (visual – tactile) stimulation paradigm was
used in right brain-damaged (RBD) patients with left
tactile extinction. Extinction is a clinical sign whereby
patients are able to detect a single stimulus presented
either to the ipsi- or contralesional side of the body, but
fail to report the contralesional stimulus when a con-
current stimulus is presented on the ipsilesional side.
Journal of Cognitive Neuroscience 14:7, pp. 1030 – 1043
Although extinction phenomena have long been studied
within different sensory modalities (vision, touch, audi-
tion, and olfaction), they also occur between sensory
modalities (Mattingley, Driver, Beschin, & Robertson,
1997). Most interestingly, cross-modal studies in RBD
patients with tactile extinction showed that visual stimuli
presented near (5 cm) the patient’s ipsilesional hand
(or cheek) produce a very strong extinction of concur-
rent tactile stimuli delivered on the contralesional hand
(or cheek). In contrast, visual stimuli presented far
(35 cm) from the patients ipsilesional hand (or cheek)
only mildly extinguish tactile stimuli simultaneously
delivered on the contralesional side of the body
(Làdavas, di Pellegrino, et al., 1998; Làdavas, Zeloni,
et al., 1998; di Pellegrino, Làdavas, et al., 1997).
This pattern of results is what should be expected if an
ipsilesional visual stimulus presented near the body
were processed by an integrated visual – tactile system
coding peripersonal space, functionally analogous to the
one described in monkeys. Due to this sensory integra-
tion, a visual stimulus presented near ipsilesional body
parts strongly activates the corresponding somatosen-
sory representation of those body parts, thus extinguish-
ing the contralesional tactile stimulation.
While the existence of a visual peripersonal space in
humans has been well documented by group studies,
only preliminary evidence based on a single-case study is
available to support the existence of a similar integrated
system coding auditory peripersonal space (Làdavas,
Pavani, & Farnè, 2001). In other words, the question
to be addressed now is whether the space immediately
surrounding our body can be activated by an auditory
stimulus. This would be possible if the human brain can
process both tactile and auditory information in an
integrated auditory –tactile system.
Neurophysiological results provide relevant clues
related to this question. Recently, Graziano, Reis, and
Gross (1999) documented neurons in monkey’s ventral
premotor cortex with tactile RFs covering the whole
contralateral side of the animal’s head. Most of these
neurons were multimodal. They were strongly activated
by tactile stimuli and also by visual and acoustic stimuli
presented close to the head, for example, within the
extension of auditory RFs whose location matched the
location of tactile RFs. Importantly, neuronal responses
almost disappeared when auditory stimuli were pre-
sented at longer distances, that is, 30 or 50 cm away
from the head. Another interesting functional property
of these neurons was that they could be strongly acti-
vated by complex sounds (e.g., burst of white noise,
jingling keys, claps, or crinkling paper), whereas pure
tones of various frequencies (200 – 3200 Hz) did not
elicit any response. Due to their functional character-
istics, these neurons may constitute the neurophysio-
logical substrate of an integrated auditory– tactile system
coding the peripersonal space near the head, that is the
peri-head space.
a b
Figure 1. Experiment 1: Schematic drawing of experimental setup.
(a) Location of tactile stimuli (indicated by arrows) for the assessment
of tactile extinction at the level of the neck. (b) The four possible
loudspeaker positions relative to patient’s head (near and far position
from right front and back spaces). Tactile stimulus location on the left
side of the neck is also indicated by an arrow. Two types of sound
(white noise and pure tone) could be delivered from each position
(sound pressure level constantly held at 70 dB).
Here, we verified whether a similar integrated system
exists in humans by investigating cross-modal extinction
between a sound and a touch in a group of 18 RBD
patients with left tactile extinction. The existence of such
system would predict that an ipsilesional auditory stim-
ulus should extinguish a contralesional tactile stimulus
(cross-modal auditory – tactile extinction). In particular,
this phenomenon should be evident when an auditory
stimulus is presented near the ipsilesional side of the
head and a tactile stimulus is simultaneously delivered to
the contralesional side of the head. This is because, due
to the activity of this integrated system, an acoustic
stimulus close to the head would activate the corre-
sponding somatosensory representation of that side of
the head. The simultaneous activation of the somato-
sensory representation of the left side of the head (by a
tactile stimulus) and of the right side of the head (by an
acoustic stimulus) should result in an extinction of those
stimuli presented in the weaker representation, that is,
that corresponding to the contralesional side of the
head (Duncan, 1996; Ward, Goodrich, & Driver, 1994).
In contrast, a significant reduction of cross-modal extinc-
tion should be expected when the auditory stimulus is
presented in the ipsilesional side, but far from the head.
This is because an auditory stimulus presented at greater
distance should activate the system encoding periper-
sonal space to a lesser degree.
Moreover, on the basis of the neurophysiological
findings (Graziano et al., 1999) auditory– tactile extinc-
tion was expected to occur in the whole space immedi-
ately surrounding the head. This is because the
topographical distribution of auditory – tactile RFs has
been shown to include both the front and the back
spaces of the animal’s head. For this reason, we inves-
tigated cross-modal effects with sound sources located
either in the front or in the back space of the patients’
head (see Figure 1).
Farnè and Làdavas 1031
 Finally, cross-modal extinction was expected to be
modulated by the complexity of the acoustic stimulus.
Since the integrated system described in monkeys by
Graziano et al. (1999) was not sensitive to simple sounds
(i.e., pure tones), a strong cross-modal extinction was
expected following the presentation of a complex
sound, like a white noise burst, but not following the
presentation of a pure tone.
These questions were investigated in 18 RBD patients
who participated to the first experiment.
EXPERIMENT 1
Results
Patients almost never produced false alarms in the
no-stimulation catch trials, and they never reported
spurious ‘‘both’’ responses relative to unilateral stimu-
lation. A one-way analysis of variance (ANOVA) was
performed on the arc sin-transformed percentage of
accuracy obtained in the unimodal tactile condition,
with type of stimulation (unilateral, bilateral) as within-
subject factor. This source of variance was highly sig-
nificant, F(1,17) = 71.86, p < .0001. Patients performed
near ceiling on trials consisting of unilateral left stim-
ulation (91% of correct responses) showing that their
tactile sensitivity was well preserved for single-stimulus
detection on the left side of the neck. In contrast,
patients only reported a small proportion of left stimuli
under bilateral stimulation (36%, p < .0001) showing
that they were affected by a severe form of unimodal
tactile extinction (see Figure 2).
To verify whether a right acoustic stimulus may
extinguish a left tactile stimulus, the amount of contrale-
Unimodal tactile extinction
Left tactile detection (% accuracy)
100
90
80
70
60
50
40
30
20
10
0 lation (unimodal tactile, cross-modal near, cross-modal
Unilateral tactile Bilateral tactile
Figure 2. The mean percentage of left contralesional detection is
reported (bars show standard error) as a function of unilateral and
bilateral stimulation in the unimodal tactile condition.
1032 Journal of Cognitive Neuroscience
sional detection obtained under bilateral cross-modal
stimulation, that is, right sound and left touch, was
compared to that obtained under unilateral left tactile
stimulation. Significantly fewer contralesional detection
in bilateral trials compared to unilateral trials indicates
cross-modal auditory –tactile extinction.
A repeated measure ANOVA with sound (white noise,
pure tone), space (front, back), and type of stimulation
(unilateral left tactile stimulation, bilateral cross-modal
stimulation in near and far position) as within-subject
factors revealed a significant main effect of sound,
F(1,17) = 54.72, p < .0001, with pure tones being
associated to more correct responses (77%) than white
noise (67%). The main factor type of stimulation was
also significant, F(2,34) = 47.02, p < .0001. Newman –
Keuls post hoc analysis showed that patients were
significantly more accurate in detecting left tactile
stimuli when presented alone (92.3%) than on bilateral
cross-modal presentations (near 54%; far 70%, p < .0001
in both comparisons). Most important, cross-modal
extinction varied according to the sound distance, being
more pronounced when the sound source was near
the patients’ head (54%) than when it was far from
it (70%, p < .003).
The interaction Sound  Space  Type of stimulation
was also significant, F(2,34) = 5.02, p < .01. As shown in
Figure 3A, when the sound was a pure tone, cross-modal
extinction was not modulated by the proximity of the
sound source to the patient’s head in the front space
(near 68% vs. far 72%, ns). Instead, the modulation was
evident in back space (near 61% vs. far 76%, p < .0001).
In contrast, when the sound was a burst of white noise
(see Figure 3B), the specificity of cross-modal extinction
for the near compared to the far space was evident both
in the front (46% vs. 69%, p < .0002) and in the back
spaces (41% vs. 63%, p < .0002).
The interaction also revealed that the back space was
more sensitive to cross-modal effects than the front
space. As can be seen in Figure 3B, cross-modal extinc-
tion induced by white noise was more severe in the back
than in the front space, both in the near (back 41% vs.
front 46%, p < .04) and in the far space (back 63% vs.
front 69%, p < .02). A similar pattern of results was
present with pure tones (Figure 3A), but only for the
near space (back 61% vs. front 68%, p < .001).
To investigate the severity of cross-modal extinction,
the amount of contralesional detection obtained under
bilateral cross-modal stimulation (i.e., right sound and
left touch) has been compared to that obtained under
bilateral unimodal stimulation (i.e., right touch and left
touch). A further one-way ANOVA with bilateral stimu-
far) as within-subject factor was separately performed for
each sound (white noise and pure tone) and space of
presentation (front and back).
The analyses showed that unimodal tactile extinction
(36% of correct responses) did not significantly differ
Volume 14, Number 7
Figure 3. The mean percen-
tage of left contralesional A
detection is reported (bars Auditory-Tactile cross-modal extinction
show standard error) as a func-

Left tactile detection (% accuracy)

tion of unilateral and bilateral 100
stimulation in the cross-modal
auditory – tactile conditions with 90
sounds presented in the near
and far locations both in the 80
front and back spaces. (A)
Accuracy following presentation 70
of pure tones. (B) Accuracy
following presentation of white 60
noise bursts. Note that accuracy
in unilateral tactile stimulation 50
is separately reported from the
condition in which sounds were
40
presented in the front (F) or
30
back (B) space.
Unilateral Unilateral Bilateral Bilateral Bilateral Bilateral
tactile (F) tactile (B) Front Near Front Far Back Near Back Far

B Auditory-Tactile cross-modal extinction

Left tactile detection (% accuracy)

100
90
80
70
60
50
40
30
Unilateral Unilateral Bilateral Bilateral Bilateral Bilateral
tactile (F) tactile (B) Front Near Front Far Back Near Back Far

from cross-modal extinction when white noise bursts Since in the preliminary auditory testing session
were presented near the patient’s head, either in the (see Methods) half of the patients failed the front/ back
front (46%) or in the back space (41%, p > .05 in both discrimination of sound source location, the differential
comparisons). In other words, the degree of interfer- auditory localization abilities of patients could have
ence exerted on the contralesional tactile perception by somewhat affected the spatial distribution of cross-
a touch or a burst of white noise was comparable. modal extinction. In order to verify this possibility, the
When white noise bursts were presented far from the same set of experimental data were also analyzed by a
patient’s head however, unimodal tactile extinction separate ANOVA in which the performance to the front/
(36%) was stronger than cross-modal extinction, both back test (correct, incorrect) was examined as between-
in the front (69%, p < .0001) and in the back spaces subjects factor, with the same within-subject factors as
(63%, p < .0005). the previous analysis. The latter analysis, besides repli-
In stark contrast, unimodal tactile extinction (36% cating the previous results, showed that this was not the
correct) was more severe than cross-modal extinction case because the factor front/back test was not signifi-
induced by pure tones presented near patients’ head, cant, nor included in any significant interaction.
both in the front (68% correct, p < .001) and in the
back spaces (60% correct, p < .001). Similarly, unimodal
Discussion
tactile extinction (36% correct) was stronger than
cross-modal extinction when pure tones were presented The results of Experiment 1 showed that in RBD patients
far from patients’ head, either in the front (72% correct, with left tactile extinction at the level of the neck, an
p < .0001) or in the back space (76% correct, p < .001). acoustic stimulus presented near the right side of the

Farnè and Làdavas 1033

head strongly interfered with the processing of a tactile
stimulus simultaneously presented on the left side of the
neck (cross-modal auditory – tactile extinction). By con-
trast, a significantly weaker interference of audition on
touch perception was observed when the same acoustic
stimulus was presented far from the patient’s right side
of the head. The differential effects produced by acous-
tic stimulation in the near versus far space were
obtained despite the fact that sound’s salience, at least
in terms of sound pressure level, was the same (70 dB)
independent of the distance of the sound source. This
finding clearly points to the processing of other auditory
cues that may be relevant for the coding of distance in
audition (see note 1).
This finding clearly shows that cross-modal auditory –
tactile extinction was mainly segregated into the peri-
personal space surrounding the head. Moreover, the
effect was most consistent when a complex sound
was presented. In particular, the spatial specificity of
auditory –tactile extinction for the peri-head space was
evident both in the front and back spaces when the
acoustic stimulus was constituted by a white noise
burst. In contrast, when the acoustic stimulus was
constituted by a pure tone the specificity for peri-head
space was present only in the back space. Thus, the
back space appeared to be more sensitive than the front
space in mediating peripersonal cross-modal effects, as
shown by the fact that both sound types induced the
strongest auditory – tactile extinction when presented
behind the head. These findings cannot be solely
explained within the theoretical framework of an inte-
grated competition between a weak representation of
the contralesional tactile input and an intact represen-
tation of the ipsilesional auditory stimulus. Models of
selective attention (Driver, Mattingley, Borden, & Davis,
1997; Duncan & Humphreys, 1989; Duncan, 1980) have
suggested that unimodal extinction is the product
of unequal strength between competitive weights,
assigned to ipsi- and contralesional stimuli, for accessing
to a limited pool of attentional resources (Driver, 1998;
di Pellegrino & De Renzi, 1995). As we outlined in relation
to visuo-tactile effects (Farnè, Pavani, Meneghello, &
Làdavas, 2000), an account based on the integrated
competition model would predict that the unbalance
caused by a unilateral lesion within a given modality
(e.g., touch) would affect other modalities (e.g., audi-
tion) to a lesser degree, because the competitive
advantage spreads to intact areas that are intercon-
nected to the damaged one. By referring to the results
of the present study, this competitive mechanism can
well explain the weak cross-modal extinction observed
when a contralesional tactile stimulus was delivered
concurrently with an ipsilesional auditory stimulus,
presented in the far-peripersonal space (see also
Làdavas et al., 2001). However, the marked cross-modal
extinction obtained by presenting an ipsilesional audi-
tory stimulus in the near-peripersonal space of the head
1034 Journal of Cognitive Neuroscience
a b
Figure 4. Experiment 2: Schematic drawing of the experimental
setup. (a) The two possible loudspeaker positions relative to patient’s
head (near and far positions from left back space). Tactile stimulus
location on the right side of the neck is also indicated by an arrow.
Only white noise bursts could be delivered from each position. (b) The
two mirror positions included in Experiment 1 are reported for
comparative purposes.
is not predicted by the integrated competition model.
Here, the degree of auditory – tactile extinction that
followed the presentation of white noise burst in the
near-peripersonal space was actually comparable to the
amount of extinction produced by bilateral unimodal
tactile stimulation. Thus, a model of integrated competi-
tion between left- and right-sided spatial representations
cannot explain the finding of the differential cross-modal
extinction found in the present study as a function of
distance from the head (near vs. far), sound complexity
(white noise vs. pure tone), or space of stimulation
(back vs. front). Therefore, the integrated competition
model (Duncan, 1996) can account only for more gen-
eral aspects of cross-modal effects, for example the
reduced, but significant amount of extinction observed
in the far-peripersonal space.
As another consequence of the unbalanced competi-
tion produced by the brain lesion, the presentation of a
stimulus into the impaired contralesional space in a
given modality evokes a weak and slowly rising activity
(Rorden, Mattingley, Karnath, & Driver, 1997). This
implies that left cross-modal extinction after right brain
damage can be found when the contralesional space
representation in one modality has been impaired by the
lesion and the ipsilesional space representation is intact.
As a corollary, left cross-modal extinction should not be
evident when both the contra- and the ipsilesional space
representations are intact.
In order to test this issue, a second investigation was
conducted in which the spatial arrangement of cross-
modal stimulation was mirror-reversed (see Figure 4).
Eight out of 18 patients took part to both investigations.
None of them manifested left neglect or contralesional
extinction in the auditory modality (see Methods). The
second experiment was constituted by two additional
conditions (run at the same time as conditions of Experi-
ment 1) in which a tactile stimulus was applied to the
right side of the neck and a left auditory stimulus could
be applied near the head (a) or far from the head (b).
In this experiment, only white noise bursts were used
as acoustic stimuli and they were presented only in the
Volume 14, Number 7
back space (Figure 4). This is because a preliminary the near and far space, thus confirming that the auditory
inspection of the results obtained in 10 patients showed modality had been spared by the lesion. However, a
that the most severe cross-modal extinction emerged certain reduction in accuracy was found when a tactile
when a white noise was presented in the back space. stimulus was simultaneously presented on the right
Therefore, this combination of sound and space of stim- ipsilesional side of their neck (Figure 5B). In the latter
ulation was the most likely to reveal the presence of a case, error consisted of incorrect ‘‘right’’ responses
mirror-reversed tactile– auditory cross-modal extinction. instead of the correct ‘‘both’’ response, that is, patients
extinguished the contralesional auditory stimulus.
EXPERIMENT 2 To verify the presence and amount of cross-modal
effects as a function of the side of acoustical stimulation,
Results patients’ performance (arcsin values of accuracy) was
Patients performed without error when sounds were submitted to a three-way ANOVA with side of sound
presented singly from the left side of their head, both in (right, left), location of sound (near, far), and type of

Figure 5. The mean percen-

tage of left contralesional A
detection is reported (bars Auditory-Tactile cross-modal extinction
show standard error) as a func-
tion of unilateral and bilateral
Left tactile detection (% accuracy)

stimulation in the cross-modal

conditions with sounds 100
presented in the near and far
locations in the back space. 90
(A) Tactile accuracy following
presentation of white noise 80
from the right. (B) Acoustic
accuracy following presentation 70
of white noise from the left.
Note that accuracy in unilateral
60
tactile stimulation is separately
reported from the condition in
which sounds were presented 50
in the near (N) or far (F)
location. 40

30
Unilateral Unilateral Bilateral Bilateral
Touch (N) Touch (F) S-T Near S-T Far

B
Tactile-Auditory cross-modal extinction
Left acoustic detection (% accuracy)

100

90

80

70

60

50

40

30
Unilateral Unilateral Bilateral Bilateral
Sound Near Sound Far T-S Near T-S Far

Farnè and Làdavas 1035

stimulation (unilateral, bilateral) as within-subject fac-
tors. The main effect side of sound was significant,
F(1,7) = 38.73, p < .0005, showing that patients’
accuracy was overall better when sounds were located
in the left (96%) as compared to the right side of their
head (71%). Since the interaction involving the three
factors was significant, F(1,7) = 7.42, p < .03, we will
focus on the description of this highest source of
variance. When sounds were presented in the right
space, the analysis performed on this subset of data
exactly replicated the results obtained in the larger
group examined in Experiment 1. Newman – Keuls test
showed that patients were more accurate in detecting
left touches under unilateral presentation (88% mean
accuracy) than under bilateral cross-modal presentation
(near 42%; far 65%, p < .002 in both comparisons). Most
important, cross-modal extinction again varied accord-
ing to sound distance (see Figure 5A), being stronger
near patients’ head (42%) than far from it (65%, p <
.005). In contrast, when sounds were presented from
the left space, only a general effect of cross-modal
extinction was found. Patients were more accurate in
reporting left sounds under unilateral presentation
(100% mean accuracy) than under bilateral cross-modal
presentation (near 93%; far 92%, p < .03 in both
comparisons). As can be clearly seen in Figure 5B,
however, no sign of spatial selectivity was found with
respect to the sound distance from patients’ head.
Finally, the severity of cross-modal extinction was
much stronger following presentation of sounds in the
right than in the left space, both near (right 42% vs. left
93%, p < .0005) and far from the head (right 65% vs. left
92%, p < .002).
Discussion
When the right acoustic stimulus and the left tactile
stimulus were competing for processing resources, the
results of the present experiment replicated those found
in Experiment 1. Auditory events strongly interfered
with left tactile stimuli when they were presented in
the peri-head space, and the competition was much
reduced when the auditory events were presented in
the far space. In contrast, when the competition was
between a right tactile stimulus and a left acoustic
stimulus, the ipsilesional stimulus induced only a weak,
although significant extinction of the contralesional
contender. In addition, the amount of competition was
comparable in the near and far space.
This finding is in accordance with a modular organ-
ization of space, which predicts a redundancy of sepa-
rate spatial maps coding space with the competition
operating among topographical maps that are function-
ally linked. The unbalance produced by a unilateral
lesion within a given modality (e.g., touch) only mildly
affects other modalities that are not impaired (e.g.,
audition), probably because the ipsilesional competitive
1036 Journal of Cognitive Neuroscience
advantage is low when competing with an intact con-
tralesional representation. In these patients, the right
tactile representation is competing with the left auditory
representation (spared by the lesion) and, as a conse-
quence, the ipsilesional stimulus induces a much weaker
interference on contralesional perception than that
exerted upon an impaired representation. In other
words, the possibility of unveiling strong cross-modal
effects seem to depend upon the presence of a clear
competitive bias favoring the ipsilesional modality.
In the same line of reasoning, the occurrence of peri-
head left-sound extinction following right-touch stimu-
lation might be found in patients whose peripersonal
auditory representation has been affected by the lesion.
Our findings suggest that such a deficit might be pro-
duced by a lesion involving, or disconnecting, the brain
structures contributing to the representation of auditory
peripersonal space. Instead, the functional integrity of at
least part of these areas would be necessary for coding
peripersonal auditory space and for inducing spatial
selective cross-modal effects of audition on touch per-
ception, as found in the present study. It would be
interesting to relate the presence and the degree of
selective auditory –tactile extinction for the near-head
space with the lesion location in the population of RBD
patients investigated. However, due to the large size and
variable extent of the lesion in our group of patients, it is
difficult to precisely relate their cross-modal behavioral
phenomena with a defined cerebral location. Neuro-
physiological studies in monkeys and fMRI evidence in
humans (Bremmer et al., 2001) have indicated that the
cerebral structures responsible of multimodal process-
ing of the peripersonal space include the frontal pre-
motor areas, parietal areas (in particular VIP), and the
putamen. As suggested above (see also Làdavas,
di Pellegrino, et al., 1998), these multimodal areas
should be (at least partially) spared by the lesion to
obtain a clear segregation of cross-modal effects near the
head. In agreement with this prediction, the most
selective cross-modal effect found in the present inves-
tigation (44% of near vs. far difference, averaged across
front and back spaces) was shown by patient B. O.,
whose lesion involved the parieto-temporal region,
whereas the least selective effect (0% of difference)
was shown by patient C. V., whose lesion affected frontal
as well as parietal and temporal areas. A similar tendency
emerged when considering data on a group basis:
Patients showing strong selective effects for the near
space (30% of near – far difference) were less fre-
quently affected by fronto- parietal lesions (33%) than
patients with less selective effects (42%). However, we
would like to make clear that these data are only
preliminary, and more detailed analyses of the extent
and precise locus of the damage are needed.
In this respect, recent ERP studies have shown that
multisensory interactions can occur early in the cortical
processing hierarchy, in putative unimodal brain regions
Volume 14, Number 7
Table 1. Demographic and Clinical Details for the Two Subgroups of Patients That Satisfied or Not (Yes, No) the Following
Criterion: When Cross-modal Extinction Produced by White Noise Burst (WN) Was Stronger in the Near versus Far Space by 30% or
More (Averaged Across Front and Back Space Stimulation) Patients Were Considered as Showing a Clear Spatial-Specific Effect
D  30% WN Sex (Female/Male) Age
Yes (n = 6) 2/4 69
No (n = 12) 5/7 62
The proportional incidence of visual neglect associated to unimodal tactile extinction is also reported.
(Foxe et al., 2000). By specifically manipulating spatial
attention, several behavioral and ERP studies have shown
that cross-modal links among vision, audition, and touch
can affect sensory perceptual processes within modality-
specific cortical regions (for a review, see Eimer & Driver,
2001). These findings point to the possible (non mutually
exclusive) existence of bottom-up, feed-forward integra-
tion in early sensory processing, and top-down modu-
lation of unimodal processing through backward
propagated activity of multimodal areas.
Besides the neuroanatomical considerations, it is also
important to establish whether other factors may affect
the presence and strength of spatially selective cross-
modal effects. A retrospective analysis on the patient
populations investigated in previous studies on visuo-
tactile extinction from our laboratory (unpublished
data) suggested that symptoms like visual neglect asso-
ciated to tactile extinction and a relatively short time
elapsed from the illness onset to patient’s testing may
represent negative concomitant factors to the emer-
gence of selective cross-modal effects in the near-
peripersonal space. When considering the presence of
neglect, Table 1 shows that the incidence of visual
neglect was higher (75%) in the subgroup of patients
who showed a weak spatial selectivity of cross-modal
extinction, as compared to those showing strongly
selective effects (30%). Table 1 also shows that spatial-
specific effects emerge more clearly in chronic patients.
These findings suggest that the severity of patients’
clinical condition (in terms of presence of neglect and
the acute/chronic phase of testing) should be consid-
ered as a negative factor, potentially obscuring the fine-
grained spatial distribution of cross-modal extinction.
Further studies are needed to clarify whether and how
these factors may interact.
GENERAL DISCUSSION
It has become abundantly clear that space is constructed
by computations involving considerable integration be-
tween different senses. One intriguing question is how
the brain can integrate spatial information deriving from
separate modalities, given that each modality initially
uses different coordinates to code space (e.g., vision in
retinotopic, but touch in somatotopic, etc.). One answer
to this question is that several neural structures contain
neurons that have been shown to respond to multiple
Education ( Years) Months Poststroke Neglect (%)
6 9 30
6 6 75
modalities. For example, Stein and Meredith (1993)
summarize a large number of single-cell studies on
multimodal neurons showing that cellular responses
can indeed be enhanced by multimodal stimulation at
a common location, as compared with unimodal base-
lines. Another example, which is also more relevant to
the purpose of the present study, is related to a recent
study by Graziano et al. (1999) in which they reported
neurons in monkey’s ventral premotor cortex with
tactile RFs that covered the contralateral side of the
head, including the back of the animal’s head, which
also responded to acoustic stimuli presented near the
head. Importantly, neurons’ responses almost disap-
peared when the auditory stimulus was presented 30
or 50 cm away from the head. Thus, this integrated
system is the best candidate system responsible of
coding auditory peripersonal space around the head.
The behavioral correlate of this neurophysiological
evidence has been revealed in the present study in
RBD patients affected by left tactile extinction at the
level of the neck. An acoustic stimulus presented near
the right side of the head (20 cm of distance) strongly
interfered with the processing of a tactile stimulus
simultaneously presented on the left side of the neck
(cross-modal auditory – tactile extinction). By contrast, a
much weaker interference of audition on touch percep-
tion was observed when the same acoustic stimulus was
presented far from the patient’s right side of the head
(70 cm), despite the fact that auditory stimulus intensity
at patient’s ear remained constant. This finding clearly
shows that cross-modal auditory – tactile extinction was
mainly segregated into the peripersonal space surround-
ing the head.
Our study clearly indicates that cross-modal competi-
tion between audition and touch is modulated by the
distance in peripersonal space of the sound source.
Distant acoustic events presented in the far space did
not compete with left tactile stimuli as did acoustic events
presented in the near space. Moreover, an ipsilesional
tactile stimulus only mildly extinguished a contralesional
acoustic stimulus presented either in the near or in the
far space. Thus, our findings are best explained as result-
ing from the processing activity of an integrated system
that control both auditory and tactile inputs within
peripersonal space of the head (peri-head). Due to this
integrated system, a right peri-head acoustic stimulus
strongly activates the somatosensory representation of
Farnè and Làdavas 1037
the head ipsilesionally, which then competes with the left
somatosensory representation of the head, activated by a
tactile stimulus. In patients with left tactile extinction,
competition favors the ipsilesional stimulus that, as a
result, extinguishes the contralesional stimulus whose
representation is assigned with weaker competitive
weights (Ward et al., 1994; di Pellegrino, Basso, & Frassi-
netti, 1997). In contrast, a distant acoustic stimulus
weakly activates the somatosensory representation of
the ipsilesional side. As a consequence, the competitive
bias is much reduced, as shown by the reduction of
cross-modal auditory– tactile extinction found far from
patient’s head. It is worthwhile to remember that, as for
visuo-tactile neurons coding visual peripersonal space
around the face (Graziano et al., 1995; Duhamel et al.,
1991, 1998), neurons coding auditory peri-head space
also show a gradient of firing that varies as a function of
stimulus distance. That is, the activation of some audi-
tory – tactile cells is greater at closer distances, being
reduced at longer distances. It is therefore possible that
the mild cross-modal extinction found in the far space
reflects the reduced neurons’ response to stimuli pre-
sented at longer distance from the head.
Interestingly, this pattern of results emerged mainly
with complex sounds, that is, when a white noise burst
was presented. Although pure tones produced a mild
cross-modal extinction, this effect appeared to be seg-
regated in the peripersonal space only behind patients’
head. In the front space, there was no differential cross-
modal extinction for the near versus the far space. These
findings suggest that patients could extract relevant
distance cues, despite the fact that sound pressure level
was kept identical irrespective of sound distance to
avoid salience differences between auditory stimulation.
In addition, the present results point to the possibility
that binaural distance cues can contribute to pure tone’s
localization, at least in the rear space.
Finally, the amount of peri-head cross-modal extinc-
tion obtained with white noise bursts (in both the front
and back spaces) was comparable to that obtained with
bilateral touches. In other words, the degree of peri-
personal cross-modal auditory – tactile and unimodal
tactile extinction did not differ in case of a complex
sound. In contrast, the degree of interference produced
by pure tones on contralesional tactile detection was
significantly smaller than that induced by a touch, even
when the pure tone was presented close to the head.
The different pattern of results found with white
noise and pure tone is in accordance with the neuro-
physiological observation reported by Graziano et al.
(1999) showing that pure tones were not efficacious in
activating multimodal neurons responsible for coding
auditory peripersonal space near the head. One way to
interpret the different effects of types of sound in the
peripersonal space is by considering the functional role
of ventral premotor (PMv) neurons that might encode
directly the position of the sound with respect to the
1038 Journal of Cognitive Neuroscience
body1 (Graziano et al., 1999). Since these neurons are
not activated by pure tones, the distance between near
and far tones cannot be estimated and, as a consequence,
cross-modal extinction is much less spatially specific.
Although this selective activation of multimodal neurons
in PMv can appear somehow surprising, it may result
from adaptive processes. It has been suggested that the
functional properties of monkeys’ multimodal neurons
are hardwired (Graziano et al., 1997) and the spatial
correspondence between RFs can be calibrated through
experience, perhaps within a critical period early in life
(Graziano et al., 1997; Salinas & Abbot, 1995). In the case
of auditory– tactile neurons, the obvious crucial experi-
ence through which spatial calibration could be achieved
consists in the repeated exposition to ecological sounds
that are more similar to a white noise burst than to a pure
tone. Thus, the reduced effect operated by a pure tone
might reflect a sort of impenetrability of the integrated
auditory – tactile system to a sound that has a little chance
to occur in nature.
A further important aspect of the head peripersonal
space that has been reported in monkeys is that tactile
RFs of multimodal neurons are rarely restricted to the
back of the head and instead include part of the cheek,
eyebrows, snout, and jaw, thus forming a complete
representation of the space around the head (Graziano
et al., 1999). In agreement with these neurophysiological
findings, the peri-head selectivity of cross-modal extinc-
tion following presentation of white noise bursts was
found both in the front and in the back spaces. How-
ever, we also found that peri-head cross-modal extinc-
tion induced by white noise was stronger in the back
space, thus suggesting that the back space may play a
major role in mediating cross-modal integration of audi-
tion and touch. This possibility is further supported by
the fact that some selectivity for the peri-head space was
also evident when pure tones were used, but only if they
were presented in the back space. Therefore, we suggest
that the latter space might represent the ideal space to
reveal auditory – tactile interactions. This better sen-
sitivity could emerge during the adaptation processes
because an object approaching the head from the back
space can be perceived only through the noise it pro-
duces, in sharp contrast with an object approaching the
head from the front space that, in normal situations,
would also benefit of visual information.
The cross-modal effects described in the present
study were found while patients were blindfolded and,
thus, did not have any visual cue about the spatial
source of auditory stimuli. This suggests that the audi-
tory peripersonal space can be coded by an integrated
system that controls directly auditory and tactile stimuli,
and that visual inputs are not necessary for achieving
bimodal auditory – tactile integration. Finally, the results
of the second experiment further demonstrated that
cross-modal auditory –tactile effects in the peripersonal
space were not dependent upon a general competition
Volume 14, Number 7
between left and right spatial representations. When the
spatial arrangement of tactile and auditory stimuli was
mirror-reversed, a small but significant amount of tac-
tile – auditory extinction was indeed found, but this was
absolutely nonselective for the space near the head.
Our findings shed new light on humans’ multisensory
integration occurring in the peripersonal space (Làdavas,
2002; in press) by showing the existence of an auditory
space representation surrounding the head. The finding
that multimodal integration involves not only somato-
sensory and visual inputs (Làdavas et al., 2000; Làdavas,
di Pellegrino, et al., 1998; Làdavas, Zeloni, et al., 1998),
but also somatosensory and auditory inputs has impor-
tant implications relative to the functional role played by
peripersonal space representations. As far as the visual –
tactile input is concerned, converging evidence from
both animal and human studies suggested that their
integrated processing in the peripersonal space would
allow to detect and localize an object approaching the
head (or the hand), before the contact with the skin
occurs, and to prepare an adequate response to it
(Graziano & Gross, 1998; Rizzolatti, Luppino, & Matelli,
1998). Depending upon its nature, the object could be
either avoided or reached and grasped.
In some circumstances, however, objects moving
towards the body are not visible, as in the dark or when
they come from the back. In these cases, the presence of
an object can be detected only by the noise it may
produce in approaching the body. On the basis of the
results of the present study, we suggest that the auditory
peripersonal space representation could allow detection
and localization of nonvisible objects approaching the
head through the noise they produce, before they reach
the body. This, in turn, would help preparing an appro-
priate orienting or avoiding reaction in response to
these objects. This function would be especially relevant
in case of objects approaching the head from the back,
since they are always outside visual control.
METHODS
Subjects
Eighteen patients with contralesional tactile extinction
were tested in the present study. Characteristics of
patients are outlined in Table 2. All of them partici-
pated in Experiment 1 (see below). A subgroup of 8
patients also took part in Experiment 2. All patients
suffered a right-hemisphere stroke, as determined by

Table 2. Clinical and Demographic Details of RBD Patients

Education Months Visual Auditory Auditory

Patient Sex Age ( Years) Poststroke Lesion Site Neglect Neglect Extinction
C. A. female 83 5 19 F, T, P no no –
C. E. male 51 8 10 F, Ic, Wm no no –
T. M. male 84 0 >24 P, O, Wm no no –
G. N. male 73 5 12 T, O, Wm yes no –
D. M. female 48 5 12 T, P yes no –
P. P. male 79 5 1 BG yes no –
L. M. G. female 60 3 1 BG yes no –
G. S. male 82 3 1 P, Ic yes no –
P. A. male 53 18 3 P, O, Ic no no no
R. M. F. female 52 8 >24 T, P no no no
B. O. male 52 5 3 P, T, Wm no no no*
R. E. female 77 5 3 F, T, P yes no no*
F. F. male 65 5 2 T, P yes no no*
G. G. female 50 8 3 F, T, P no no no*
D. L. female 56 5 2 F, T, P no no no*
T. O. male 64 5 2 F, T, P, Ic yes no no*
C. V. male 69 14 4 F, T, P yes no no*
B. L. male 65 5 1 F, T, P no no no*

‘‘Lesion site’’ column indicates which structures were involved by the lesion. F = frontal; T = temporal, P = parietal; O = occipital; Ic = internal
capsule; BG = basal ganglia; Wm = white matter. The table also shows that patients who participated to the second experiment (indicated by an
asterisk) were not affected by auditory extinction (see Methods).

Farnè and Làdavas 1039

cranial CT scan. Nine of them showed signs of visual
neglect.
Patients were selected according to the absence of
acoustical sensory loss and right – left ear difference
(>10 dB) in hearing, as assessed by an audiometry test.
Moreover, patients did not show any sign of auditory
neglect, when tested with the same auditory stimuli
employed in the experimental investigation (white noise
bursts and 1.5 kHz pure tones). When 20 unilateral left
and right acoustic stimuli were resented, patients’ per-
formance was nearly perfect.
Patients were also selected according to the absence
of obvious tactile sensory loss. Tactile stimuli were
delivered behind the ears on the lateral surface of the
neck, just below the mastoid process of the temporal
bone. In order to be included in the study, patients had
to show a good performance (80% correct) in detect-
ing single left and right tactile stimulation, showing in
this way a well-preserved tactile sensitivity in relation to
the neck. Most important, patients were selected accord-
ing to the presence of left tactile extinction (left right
difference 20%) under the condition of double simul-
taneous stimulation applied at the level of the neck.
Patients who also participated in Experiment 2 were
submitted to an additional evaluation of unimodal audi-
tory extinction, as assessed by using the same sounds
employed for the experimental investigation. When 20
unilateral left and right acoustic stimuli were presented
either unilaterally or bilaterally at ear level, the perform-
ance of each patient was almost errorless, thus showing
no sign of auditory extinction.
Materials and Procedure
The apparatus for presentation of stimuli consisted of
a loudspeaker (0.3 W, 8 ), a touch-sensitive sensor
(1 cm2), and a remote switch that was connected to an
optically isolated electric circuit. Two positions of the
switch (pressed = on; released = off ) controlled sound
emission through the loudspeaker. With the switch set
in the ‘‘on’’ position, the contact of the sensor with
the skin triggered, besides the tactile stimulation, the
simultaneous production of a sound whose duration
was identical to the time of contact between the sensor
and the skin. Two types of sounds could be delivered,
either wideband white noise bursts or pure tones
(1.5 kHz). Tactile stimuli were brief (<1 sec) light con-
tacts of the sensor with the patient’s skin. Tactile stimuli
were applied on the lateral surface of the neck, behind
the ear, and just below the mastoid process of the
temporal bone.
Experiment 1
Each patient sat blindfolded in a silent room and was
instructed to keep the head straight throughout the
entire experiment. The experimenter sat behind the
1040 Journal of Cognitive Neuroscience
patient, with the sensor firmly attached to the index
finger of her/his left hand, and checked the patient’s
head position before each trial. The loudspeaker was
mounted at ear level on an adjustable metal stand and it
was positioned on the right of the patient’s head, either
in front or behind it (see Figure 1b). Specifically, the
loudspeaker was silently placed in one of four locations
along a line between sagittal and coronal planes: (1) in
the front right quadrant 20 cm from the patient’s head;
(2) in the front right quadrant 70 cm from the patient’s
head; (3) in the back right quadrant 20 cm from the
patient’s head; (4) in the back right quadrant 70 cm from
the patient’s head. Sound intensity was identical for
each loudspeaker position and type of sound. Sound
pressure level, as measured at the patient’s ear was set at
70 dB by adjusting sound intensity by means of a sound
meter.
The angular position of the loudspeaker was the same
in both the front and the back spaces, and it was
determined for each subject in order to fall outside
her/his cone of acoustical confusion (Moore, 1995;
Moore & King, 1999). To this aim, subjects were sub-
mitted to a front/back discrimination task in which they
were required to localize the source of singly emitted
sounds. Two identical loudspeakers were initially
located at 458 from the sagittal plane in both spaces. A
sound (either a white noise or a pure tone, assessed in
separate blocks) was randomly delivered through one of
the loudspeakers and the subject was required to ver-
bally judge whether it came from the front or the rear
space. In case of correct performance (80% correct on
10 trials per each type of sound and distance), the
experiment was successively run with the loudspeaker
located at 458 from the sagittal plane in both quadrants.
Otherwise (80% correct), both loudspeakers were
displaced by 158 toward the sagittal plane, and the test
was repeated. If the patient was still unable to correctly
locate sounds in the front or in the back space after two
displacements (i.e., at 158 from the sagittal plane), the
experiment was run with the loudspeaker along the
458 line between sagittal and coronal axes. Half of the
subjects correctly performed the task with a 458 posi-
tioning whereas the remaining half was still incorrect
after 308 of displacement. Therefore, the angular posi-
tion of the loudspeaker during the experimental inves-
tigation was 458 for all the subjects. As it has been
reported for normal people (Brungart, 1999), near/far
discrimination appeared to be a very difficult task and
was not assessed systematically in our group of patients;
a few patients, among those who performed correctly
the front/ back discrimination test, failed to judge
whether a sound came from the near or far location
when submitted to a near/far discrimination test.
Experiment 1 consisted of nine different conditions:
a unimodal tactile condition and eight cross-modal
auditory – tactile conditions, one for each combination
of loudspeaker location (front and back spaces),
Volume 14, Number 7
distance of the loudspeaker from the patient’s head
(near and far), and type of sound (white noise and pure
tone). In the unimodal tactile condition, the experi-
menter had a touch-sensitive sensor attached to the
index finger of both hands. In this condition, the switch
was in the ‘‘off’’ position, that is, sensors did not trigger
any sound and they were used to obtain identical tactile
stimulation on both sides of the neck. Four types of
trials were presented: (1) a single touch on the left; (2) a
single touch on the right; (3) two simultaneous touches,
one on the left and the other on the right; (4) no tactile
stimulation (catch trials). Patients were informed they
would feel a light touch on the left or the right side of
the neck or on both sides simultaneously. They were
also informed that occasionally they would feel nothing
at all. Before each stimulation, they were verbally
prompted by the experimenter to summon attention
to the next upcoming event. In cross-modal conditions,
the remote switch was held by the experimenter in
her/his right hand to control sound emission. Four
types of trials were presented: (1) a single touch on
the left; (2) a single sound on the right (obtained by a
contact between the sensor and the experimenter’s
thumb); (3) a touch on the left and a simultaneous
sound on the right; (4) no stimulation (catch trials).
Patients were informed they would feel a light touch on
the left side of the neck or hear a sound on the right
side of the head or a touch on the left side and a
simultaneous sound on the right side while, occasion-
ally, they would neither feel nor hear anything.
In all conditions, patients were asked to verbally
report what they perceived by saying ‘‘left,’’ ‘‘right,’’
‘‘both,’’ or ‘‘none’’ irrespective of the sensory modality,
immediately after the stimulation. Whenever they failed
to respond spontaneously, they were prompted by the
experimenter. A second experimenter recorded the
response after each trial. Each condition was composed
by 40 trials presented in the same pseudorandom order
in each condition: 10 single left stimulation, 10 single
right stimulation, 10 bilateral stimulation, and 10 catch
trials. Conditions were run per block, and their order
was randomized across sessions and subjects. Patients
were submitted twice to the entire protocol, run in
separate sessions the same day and/or in two consec-
utive days.
Experiment 2
Apparatus and procedures were identical to those em-
ployed in Experiment 1, except as noted. The experi-
menter had the sensor attached to the index finger of
her/his right hand. The loudspeaker was positioned in
the left back space in one of two positions: (1) in the
back left quadrant 20 cm from patients’ head; (2) in the
back left quadrant 70 cm from patients’ head (see
Figure 4a). Each patient was assessed with the loud-
speaker located in the mirror angular positions used in
the first experiment. Experiment 2 consisted of two
cross-modal tactile– auditory conditions, one for each
distance of the loudspeaker from patients’ head. Only
white noise bursts were presented. Four types of trials
were presented: (1) a single touch on the right; (2) a
single sound on the left; (3) a touch on the right and a
simultaneous sound on the left; (4) no stimulation at all
(catch trials). Patients were informed they would feel a
light touch on the right side of the neck or hear a sound
on the left side of the head or both stimuli simulta-
neously and that, occasionally, they would feel and hear
nothing at all.
Acknowledgments
We thank all the subjects for their collaboration. We also thank
N. Allegri and C. Soncini for their help in collecting the data,
and Daniele Maurizzi for his technical support in constructing
the experimental apparatus. We also thank F. Frassinetti for
reading CT scans. This work was supported by grants from
MURST and CNR.
Reprint requests should be sent to Alessandro Farnè, Diparti-
mento di Psicologia, Università di Bologna, via le Berti Pichat,
5-40127 Bologna, Italy, or via e-mail: farne@psibo.unibo.it.
Note
1. The auditory system processes different spatial acoustic
cues, including time of arrival and level differences at each ear,
and spectral cues generated by filtering effects of head and
external ears (King, Schnupp, & Doubell, 2001; Blauert, 1983).
This computation is partly done at brainstem level (Middle-
brooks & Green, 1991), with a substantial contribution of
primary auditory cortex and parieto-prefrontal regions that
also integrate information from other sensory modalities
(Schroeder et al., 2001; Bushara et al., 1999; Weeks et al.,
1999). Many details of auditory distance perception in humans
remain obscure. When free-field, amplitude-constant sounds
are presented in closed room, as in the present study, auditory
distance is mainly coded through reverberation cues based on
the ratio between the sound energy reaching the ears directly
and via reflections on the walls (Bronkhorst & Houtgast, 1999),
especially when sounds originate within 1 m from the ears. In
this proximal region, monaural spectral cues and interaural
level differences are modified by distance (Brungart, Durlach,
& Rabinowitz, 1999; Brungart & Rabinowitz, 1999; Moore &
King, 1999) and localization performance reliably rely on
binaural distance cues, whose relevance is comparable to that
provided by amplitude cues (Brungart, 1999).
REFERENCES
Blauert, J. (1983). Spatial hearing: The psychophysics of
human sound localization. Cambridge: MIT Press.
Bremmer, F., Schlack, A., Shah, N. J., Zafiris, O., Kubischik,
M., Hoffmann, K., Zilles, K., & Fink, G. R. (2001).
Polymodal motion processing in posterior parietal and
premotor cortex: A human fMRI study strongly implies
equivalencies between humans and monkeys. Neuron, 29,
287 – 296.
Bronkhorst, A. W., & Houtgast, T. (1999). Auditory distance
perception in rooms. Nature, 397, 517 – 520.
Farnè and Làdavas 1041
Brungart, D. S. (1999). Auditory localization of nearby sources:
III. Stimulus effects. Journal of the Acoustical Society of
America, 106, 3589 – 3602.
Brungart, D. S., Durlach, N. I., & Rabinowitz, W. M. (1999).
Auditory localization of nearby sources: II. Localization of
a broadband source. Journal of the Acoustical Society
of America, 106, 1465 – 1479.
Brungart, D. S., & Rabinowitz, W. M. (1999). Auditory
localization of nearby sources. Head-related transfer
functions. Journal of the Acoustical Society of America, 106,
1956 – 1968.
Bushara, K. O., Weeks, R. A., Ishii, K., Catalan, M.-J., Tian, B.,
Rauschecker, J. P., & Hallett, M. (1999). Modality-specific
frontal and parietal areas for auditory and visual spatial
localization in humans. Nature Neuroscience, 2, 759 – 766.
di Pellegrino, G., Basso, G., & Frassinetti, F. (1997).
Spatial extinction to double asynchronous stimulation.
Neuropsychologia, 35, 1215 – 1223.
di Pellegrino, G., & De Renzi, E. (1995). An experimental
investigation on the nature of extinction. Neuropsychologia,
33, 153 – 170.
di Pellegrino, G., Làdavas, E., & Farnè, A. (1997). Seeing where
your hands are. Nature, 338, 730.
Driver, J. (1998). The neuropsychology of spatial attention. In
H. Pashler (Ed.), Attention (pp. 297 – 340). Hove: Psychology
Press.
Driver, J., Mattingley, J. B., Rorden, C., & Davis, G. (1997).
Extinction as a paradigm measure of attentional bias and
restricted capacity following brain injury. In P. Thier & H. O.
Karnath (Eds.), Parietal lobe contributions to orientation
in 3D space (pp. 401 – 429). Berlin: Springer-Verlag.
Duhamel, J. R., Colby, C. L., & Goldberg, M. E. (1991).
Congruent representation of visual and somatosensory
space in single neurons of monkey ventral intra-parietal
area (area VIP). In J. Paillard (Ed.), Brain and space
(pp. 223 – 236). Oxford: Oxford University Press.
Duhamel, J. R., Colby, C. L., & Goldberg, M. E. (1998). Ventral
intraparietal area of the macaque: Congruent visual and
somatic response properties. Journal of Neurophysiology,
79, 126 – 136.
Duncan, J. (1980). The locus of interference in the
perception of simultaneous stimuli. Psychological Review,
87, 272 – 300.
Duncan, J. (1996). Coordinated brain systems in selective
perception and action. In T. Inui & J. L. McClelland (Eds.),
Attention and performance (pp. 549 – 578). Cambridge:
MIT Press.
Duncan, J., & Humphreys, G. W. (1989). Visual search and
stimulus similarity. Psychological Review, 96, 433 – 458.
Eimer, M., & Driver, J. (2001). Crossmodal links in endogenous
and exogenous spatial attention: Evidence from
event-related brain potential studies. Neuroscience and
Biobehavioral Reviews, 25, 497 – 511.
Farnè, A., Pavani, F., Meneghello, F., & Làdavas, E. (2000). Left
tactile extinction following visual stimulation of a rubber
hand. Brain, 123, 2350 – 2360.
Fogassi, L., Gallese, V., Fadiga, L., Luppino, G., Matelli, M., &
Rizzolatti, G. (1996). Coding of peripersonal space in inferior
premotor cortex (area F4). Journal of Neurophysiology, 76,
141 – 157.
Foxe, J. J., Morocz, I. A., Murray, M. M., Higgins, B. A., Javitt,
D. C., & Schroeder, C. E. (2000). Multisensory auditory-
somatosensory interactions in early cortical processing
revealed by high-density electrical mapping. Cognitive Brain
Research, 10, 77 – 83.
Gentilucci, M., Fogassi, L., Luppino, G., Matelli, M., Camarda,
R., & Rizzolatti, G. (1988). Functional organization of inferior
area 6 in the macaque monkey: I. Somatotopy and the
1042 Journal of Cognitive Neuroscience
control of proximal movements. Experimental Brain
Research, 71, 475 – 490.
Gentilucci, M., Scandolara, C., Pigarev, I. N., & Rizzolatti, G.
(1983). Visual responses in the postarcuate cortex
(area 6) of the monkey that are independent of eye
position. Experimental Brain Research, 50, 464 – 468.
Graziano, M. S. A., & Gross, C. G. (1993). A bimodal map of
space: Tactile receptive fields in the macaque putamen with
corresponding visual receptive fields. Experimental Brain
Research, 9, 96 – 109.
Graziano, M. S., & Gross, C. G. (1995). The representation
of extrapersonal space: A possible role for bimodal,
visual – tactile neurons. In M. S. Gazzaniga (Ed.), The
cognitive neuroscience (pp. 1021 – 1034). Cambridge:
MIT Press.
Graziano, M. S., & Gross, C. G. (1998). Spatial maps for the
control of movement. Current Opinion in Neurobiology, 8,
195 – 201.
Graziano, M. S. A., Hu, X. T., & Gross, C. G. (1997). Visuospatial
properties of ventral premotor cortex. Journal of Neuro-
physiology, 77, 2268 – 2292.
Graziano, M. S. A., Reis, L. A. J., & Gross, C. G. (1999). A
neuronal representation of the location of nearby sounds.
Nature, 397, 428 – 430.
King, A. J., Schnupp J. W. H., & Doubell, T. P. (2001). The
shape of ears to come: Dynamic coding of auditory space.
Trends in Cognitive Sciences, 5, 261 – 270.
Làdavas, E. (in press). Visual peripersonal space in humans. In
H. O. Karnath, A. D. Milner, & G. Vallar (Eds.), The cognitive
and neural bases of spatial neglect. Oxford University
Press.
Làdavas, E. (2002). Functional and dynamic properties of visual
peripersonal space in humans. Trends in Cognitive Sciences,
6, 17 – 22.
Làdavas, E., di Pellegrino, G., Farnè, A., & Zeloni, G. (1998).
Neuropsychological evidence of an integrated visuotactile
representation of peripersonal space in humans. Journal of
Cognitive Neuroscience, 10, 581 – 589.
Làdavas, E., Farnè, A., Zeloni, G., & di Pellegrino, G. (2000).
Seeing or not seeing where your hands are. Experimental
Brain Research, 131, 458 – 467.
Làdavas, E., Pavani, F., & Farnè, A. (2001). Auditory periper-
sonal space in humans: A case of auditory – tactile extinction.
Neurocase, 7, 97 – 103.
Làdavas, E., Zeloni, G., & Farnè, A. (1998). Visual periper-
sonal space centred on the face in humans. Brain, 121,
2317 – 2326.
Mattingley, J., Driver, J., Beschin, N., & Robertson, I. H. (1997).
Attentional competition between modalities: Extinction
between touch and vision after right hemisphere damage.
Neuropsychologia, 35, 867 – 880.
Middlebrooks, J. C., & Green, D. M. (1991). Sound localization
by human listeners. Annual Review of Psychology, 42,
135 – 159.
Moore, B. C. J. (1995). Hearing. San Diego, CA: Academic Press.
Moore, D. R., & King, A. J. (1999). Auditory perception:
The near and far of sound localization. Current Biology, 9,
R361 – R363.
Rizzolatti, G., Luppino, G., & Matelli, M. (1998). The
organization of the cortical motor system: New concepts.
Electroencephalography and Clinical Neurophysiology,
106, 283 – 296.
Rizzolatti, G., Scandolara, C., Matelli, M., & Gentilucci, M.
(1981). Afferent properties of periarcuate neurons in
macaque monkeys: II. Visual responses. Behavioral Brain
Research, 2, 146 – 163.
Rorden, C., Mattingley, J. B., Karnath, H. O., & Driver, J.
(1997). Visual extinction and prior entry: Impaired
Volume 14, Number 7
 perception of temporal order with intact motion
perception after unilateral parietal damage. Neuropsycho-
logia, 35, 421 – 433.
Salinas, E., & Abbot, L. F. (1995). Transfer of coded information
from sensory to motor networks. Journal of Neuroscience,
15, 6461 – 6474.
Schroeder, C. E., Lindsley, R. W., Specht, C., Marcovici, A.,
Smiley, J. F., & Javitt, D. C. (2001). Somatosensory input to
auditory association cortex in the macaque monkey. Journal
of Neurophysiology, 85, 1322 – 1327.
Stein, B. E., & Meredith, M. A. (1993). The merging of the
senses. Cambridge: MIT Press.
Ward, R., Goodrich, S., & Driver, J. (1994). Grouping
reduces visual extinction: Neuropsychological evidence
for weight linkage in visual selection. Visual Cognition,
1, 101 – 129.
Weeks, R. A., Aziz-Sultan, A., Bushara, K. O., Tian, B.,
Wessinger, C. M., Dang, N., Rauschecker, J. P., & Hallett, M.
(1999). A PET study of human auditory spatial processing.
Neuroscience Letters, 262, 155 – 158.
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