European Journal of Neuroscience, Vol. 18, pp.

704±708, 2003 ß Federation of European Neuroscience Societies

Language perception activates the hand motor cortex:

implications for motor theories of speech perception

Agnes FloÈel,1,2 Tanja Ellger,2 Caterina Breitenstein2 and Stefan Knecht2

1
Human Cortical Physiology Section, National Institute of Neurological Disorders and Stroke, NIH, 10 Center Drive, Building 10,
Bethesda, MD 20892-1430, USA
2
Department of Neurology, University of MuÈnster, Germany

Keywords: gesture, mirror neurons, motor system, motor theory of speech perception, transcranial magnetic stimulation

Abstract
The precise mechanisms of how speech may have developed are still unknown to a large extent. Gestures have proven a powerful
concept for explaining how planning and analysing of motor acts could have evolved into verbal communication. According to this
concept, development of an action-perception network allowed for coding and decoding of communicative gestures. These were
manual or manual/articulatory in the beginning and then became increasingly elaborate in the articulatory mode. The theory predicts
that listening to the `gestures' that compose spoken language should activate an extended articulatory and manual action-perception
network. To examine this hypothesis, we assessed the effects of language on cortical excitability of the hand muscle representation by
transcranial magnetic stimulation. We found the hand motor system to be activated by linguistic tasks, most notably pure linguistic
perception, but not by auditory or visuospatial processing. The amount of motor system activation was comparable in both hemi-
spheres. Our data support the theory that language may have evolved within a general and bilateral action-perception network.

Introduction
Hypotheses about the evolution of spoken language rest on a narrow
basis of data (Hauser et al., 2002; Nowak et al., 2002). These data
come from studies of primate social behaviour or animal communica-
tion, developmental, genetic and anatomical investigations on lan-
guage, learning theory and evolutionary mathematics (for review see
Nowak et al., 2002). One in¯uential theory to explain the evolution of
communication has proposed that language could have evolved from
gestures (Hewes, 1973; Corballis, 1992; Rizzolatti & Arbib, 1999). In
this frame of reference, production and perception of spoken language
can be viewed as communication through phonetic gestures (`motor
theory of speech perception') (Liberman & Mattingly, 1985). If there
was such an evolutionary link between the hand motor and language
system, listening to `gestures' that compose spoken language should
activate an extended articulatory/manual action-perception network.
In the present study, we examined if, and to what extent, language
activates the hand motor system. Transcranial magnetic stimulation
was used to assess the effects of language on motor cortex excitability.
We contrasted changes in the amplitude of hand motor-evoked poten-
tials (MEPs) during language tasks with changes in MEPs during non-
linguistic visual and auditory control tasks.
Materials and methods
Human subjects
Subjects were selected from a cohort of 324 healthy volunteers
previously assessed for language dominance by functional transcranial
Correspondence: Dr Agnes FloÈel, 1Human Cortical Physiology Section, as above.
E-mail: floela@ninds.nih.gov
Received 18 February 2003, revised 5 May 2003, accepted 12 May 2003
Doppler ultrasonography during a word generation task (Knecht et al.,
2000a,b). Subjects were excluded if information from a standardized
questionnaire suggested neurological, psychiatric and medical disor-
ders. Handedness was assessed with the Edinburgh Handedness
Inventory (Old®eld, 1971). The study was approved by the local ethics
committee. Each individual gave written informed consent. Tasks and
testing procedures were in accordance with institutional guidelines and
the study conforms with `The Code of Ethics of the World Medical
Association' (Declaration of Helsinki).
Experimental tasks
In three consecutive experiments, we attempted to isolate the critical
components (articulatory linguistic, auditory, cognitive and visuospa-
tial attentional) which activate the motor system. Experiment I exam-
ined differences based on side of language dominance and handedness
and experiments II and III extended the ®ndings of experiment I to
other types of linguistic and control tasks.
Experiment I
Twenty-six subjects (12 men) were selected to assess the in¯uence of
handedness and hemispheric dominance for language on speech and
speech-related activation of the hand motor cortex (seven left-handed
left-hemispheric language dominant (LH-LD) subjects, six right-
handed right-hemispheric language dominant (RH-RD) subjects,
seven right-handed left-hemispheric dominant (RH-LD) subjects
and six left-handed right-hemispheric dominant (LH-RD) subjects).
This aimed to examine if a possible increase in motor cortex excit-
ability was restricted to the subject's language dominant hemisphere or
to the dominant hand, as proposed by a previous report (Tokimura
et al., 1996). The four groups did not signi®cantly differ in age, sex,
years of education and number of languages spoken ¯uently.

doi:10.1046/j.1460-9568.2003.02774.x

Experiments II and III
As no signi®cant effects were detected for handedness or hemispheric
dominance for language in the ®rst experiment, the second and third
experiments were conducted with RH-LD subjects only.
In experiment II, 10 subjects took part (®ve men) and 12 subjects
(six men) were included in experiment III. For age, handedness scores
and language laterality indices, see Table 1).
In the ®rst experiment, four different tasks were used: `reading
aloud' (paragraph from a physiology textbook), `reading silently' (para-
graph from a physiology textbook), speaking spontaneously (`speaking
spontan.') and producing phonemes (`lala'). These tasks were identical
to those employed in the study by Tokimura et al. (1996).
In the second experiment, we aimed to control that unspeci®c
arousal mechanisms led to the bilateral facilitation observed in the
®rst experiment. Two non-speech control tasks were administered.
Subjects had to memorize non-verbalizable abstract ®gures (`Kimura
pictures') (Kimura, 1963) and detect word pairs in which the ®rst word
ended with an e and the second started with an e (`ee'). Performance on
`Kimura pictures' was probed in a recognition test immediately after
the experimental task. Performance on `ee' was tested by having
subjects report the number of ee combinations found in the text
immediately after the experimental task `ee'. Additionally, four speech
or speech-like tasks were given to each subject: `humming' (produc-
tion of sounds without speech) and three tasks identical to experiment
1: `lala', `reading silently' and `reading aloud'.
In the third experiment, we wanted to examine if the increased
excitability during speaking and reading was merely due to speech-
related vocal and subvocal activation of articulatory muscles. A group of
12 RH-LD subjects took ®ve different tasks: three perceptive linguistic
tasks, where subjects had to listen to a well-known fairy-tale (`listening
to fairy-tales'), listen to short simple sentences (`listening to sentences')
and listen to white noise (`white noise'). The fourth and ®fth tasks were
reading a fairy-tale aloud (`reading aloud') and reading a fairy-tale
silently (`reading silently'). Linguistic processing of the sentences and
fairy-tale was ensured by probing subjects on the content of either.
Experimental protocol
Subjects were seated in a comfortable chair. Focal transcranial magnetic
stimulation with a ®gure-of-eight coil (Magstim Rapid stimulator;
Magstim, Whitland, South-west Wales, UK) over the right or left
primary motor cortex was used to elicit MEPs in the ®rst dorsal
interosseous muscle of the relaxed contralateral hand. The coil was held
tangentially on the scalp over the optimal spot for eliciting MEPs in the
contralateral ®rst dorsal interosseous muscle. Orientation was approxi-
mately perpendicular to the central sulcus, 458 from the anterior±poster-
ior axis, with the handle pointing posteriorly. The MEPs were recorded
by electromyography of the ®rst dorsal interosseous muscle, with the
active electrode placed over the motor point and the reference electrode
placed over the interphalangeal joint and a sampling rate of 1 kHz/
Table 1. Subject characteristics
Age Handedness
n (years) score
Experiment I 26 29  5.5 (26±42) RH: 90  19 (40±100)
LH: 74  22 ( 23 to
Experiment II 10 26  5.2 (19±37) 89  20 (40±100)
Experiment III 12 23  2.2 (21±28) 97  6 (80±100)
Data are presented as means  SD (and range). n, number; RH, right-handed; LH, left-handed; LD, left-hemispheric language dominant; RD, right-hemispheric
language dominant.
ß 2003 Federation of European Neuroscience Societies, European Journal of Neuroscience, 18, 704±708
Language and the motor system 705
channel. Responses were ampli®ed using a low-frequency ®lter of 20 Hz
and a high-frequency ®lter of 5 kHz (sensitivity 100 mV/D, sweep
10 ms/D). Resting motor thresholds of the right and left ®rst dorsal
interosseous muscles were assessed (Rossini et al., 1994) and the output
of the stimulator was subsequently adjusted to approximately 10±15%
above resting motor threshold to produce an electromyography response
with a peak-to-peak amplitude of about 300±500 mV for the baseline
resting condition. The subjects were required to maintain complete
muscle relaxation of the hand muscles during all tasks. Background
electromyography activity was monitored to ensure complete relaxation
of hand muscles during the study (sensitivity 50 mV/D, sweep 10 ms/D).
The MEP peak-to-peak amplitudes were analysed for each trial. Trials
were discarded from analysis if the electromyography revealed that the
task hand was not fully relaxed.
To assess hemispheric differences in MEP changes on the experi-
mental tasks, each subject was stimulated over the left and right
hemispheres on each task. The order of stimulation sites was counter-
balanced between subjects (for experimental set-up, see Fig. 1).
On each block, 32 trials were recorded: ®rst, for the baseline
condition, 16 stimuli with the subjects at complete rest (`baseline')
and the next 16 stimuli during task performance (`experimental').
Constant coil positioning was assured by ®xing the coil to the scalp by
Fig. 1. Experimental set-up. The coil is positioned over the right hemisphere
and motor-evoked potentials (MEPs) are recorded from the contralateral hand
using electromyography (EMG) either during baseline (rest) or during the
experimental task (in this case listening). Amplitudes of MEPs were calculated
automatically (Medtronic Functional Diagnostics, Keypoint, Minneapolis, MN,
USA). For each subject, EMG amplitudes were pooled across each baseline and
each experimental task.
Laterality
index
LD: 4.04  1.93 (1.62±8.19)
100) RD: 4.04  1.28 ( 7.03 to 2.80)
4.26  1.90 (2.48±8.19)
4.51  1.90 (1.73±9.23)
706 A. FloÈel et al.
Table 2. Resting motor thresholds and applied stimulus intensities in
experiments I±III
MT left FDI MT right FDI SI left FDI SI right FDI
muscle (%) muscle (%) muscle (%) muscle (%)
Experiment I 53  6.6 53  7.1 58  7.8 58  7.5
Experiment II 50  2.5 50  2.8 54  2.3 54  2.9
Experiment III 55  3.0 54  3.1 59  3.2 59  3.0

Mean (SD) percentage of maximum stimulator output. MT, motor
threshold; SI, stimulus intensity; FDI, first dorsal interosseus.
a specially designed coil-holder. The subsequent task started again
with the baseline condition. Subjects completed all tasks with the coil
over one hemisphere and MEPs elicited from the contralateral hand.
The stimulating coil was then switched to the other hemisphere. The
order of hemisphere and the order of condition were pseudorando-
mized across subjects. We used a ®xed sequence of baseline-experi-
mental conditions. To assure that no systematic error resulted, we
compared baseline conditions within each experiment for signi®cant
differences. If no signi®cant differences were found, a carry-over
effect from baseline transcranial magnetic stimulation effects to the
experimental condition would be unlikely.
Data analysis
For all experiments, ANOVAs with the repeated factors side (2), task
(experiment I, 4; experiment II, 6 and experiment III, 5) and condition
(2) were conducted. Analysis of experiment I also included the
between-subject factor group (LHLD, LH-RD, RH-LD and RH-
RD). Post-hoc analyses were conducted using Tukey tests or paired
t-tests, as appropriate.
Results
Motor threshold, stimulus intensity and electromyography
level of task hand
Resting motor thresholds and stimulus intensities were not different
between the left and right hand muscles (see Table 2). Therefore, left
versus right baseline differences in cortico-motoneuronal excitability
cannot explain possible differences of the experimental conditions.
Age
Groups I and III differed signi®cantly in age (F2,45 ˆ 7.10; Tukey
P ˆ 0.002). As the correlations between age and MEP effects were not
Fig. 2. Raw motor-evoked potentials (MEPs) of one of the subjects of experiment III (female, 22 years old, handedness score of 100, language laterality index of 4.5)
during a baseline condition (mean MEP amplitude 266 mV) and two different experimental conditions: `white noise' (mean MEP amplitude 327 mV) and `listening to
sentences' (mean MEP amplitude 474 mV). Eight MEPs are shown for each condition.
ß 2003 Federation of European Neuroscience Societies, European Journal of Neuroscience, 18, 704±708
signi®cant for any of the tasks (all r < 0.28), it was not considered
necessary to use age as a covariate in statistical analyses.
Motor-evoked potential amplitudes
There were no signi®cant differences in the number of rejected trials
between `baseline' and `experimental' conditions for each task or across
tasks. The baselines of the different experimental conditions were not
signi®cantly different within each of the three experiments. Therefore,
comparisons between baselines within each experiment will not be
reported below. Overall, the amplitude of baseline MEPs was arbitrarily
set between 300 and 500 mV by the experimenter. Therefore, baseline
MEP amplitudes cannot be compared between experiments.
The raw data for individual MEP trials are presented in Fig. 2 for
experiment III's conditions `baseline', `white noise' and `listening to
sentences'.
Experiment I (see Fig. 3a)
No signi®cant difference for side of stimulation was detected. Interaction
of condition by task (F3,66 ˆ 3.74, P ˆ 0.02) was signi®cant. The diff-
erence between baseline and experimental conditions was greatest for
`reading aloud', followed by `still reading', `producing phonemes' and
`speaking spontaneously' (all t25  4.67, all P < 0.001).
Furthermore, the main effect of task for the experimental condition
(F3,75 ˆ 4.41, P ˆ 0.007) could be explained with signi®cantly smaller
MEPs for `reading silently' as compared with `reading aloud', `lala'
and `speaking spontaneously' (all t25  2.80, P  0.001). The latter
three conditions were not signi®cantly different.
The lack of signi®cant effects involving the factors group and side
indicates that the experimental manipulation was independent of
language dominance, handedness and stimulated hemisphere.
Experiment II (see Fig. 3b)
Again, no signi®cant difference for side of stimulation was detected.
As for experiment I, the interaction of task by condition yielded
signi®cance (F5,40 ˆ 4.87, P ˆ 0.01). Post-hoc tests revealed signi®-
cantly greater MEPs during experimental compared with baseline
conditions for all four speech tasks (all t9  3.51, P  0.007), The
respective MEP difference was, however, not signi®cant for `Kimura
pictures' or for `ee'.
The main effect of task for the experimental condition (F5,45 ˆ
5.62, P < 0.001) could be explained with signi®cantly smaller MEPs
for `ee' compared with `reading silently', `reading aloud', `lala' and
`humming' and signi®cantly smaller MEPs for `Kimura pictures' as

Fig. 3. Experiment I (top): mean ( SD) motor-evoked potential (MEP)
amplitudes during the four different baselines (base) and the respective experi-
mental (exp.) conditions. spontan., spontaneously. Experiment II (middle):
mean (SD) MEP amplitudes during the three different baselines (base) and
the respective experimental (exp.) conditions. Experiment III (bottom): mean
(SD) MEP amplitudes during the three different baselines (base) and the
respective experimental (exp.) conditions.
compared with `reading aloud', `lala' and `humming' (all t9  2.90, all
P < 0.02).
Experiment III (see Fig. 3c)
As for experiments I and II, no effect for side of stimulation was
detected. The interaction of task by condition yielded signi®cance
(F4,44 ˆ 4.81, P ˆ 0.009). The main effects of condition at each level of
task showed signi®cantly greater MEPs during experimental compared
with baseline conditions for `listening to fairy-tales', `listening to
sentences', `reading aloud' and `reading silently' (all t11  3.89,
P  0.003). The respective MEP difference was not signi®cant for
`white noise'.
Furthermore, the main effect of task for the experimental condition
(F4,44 ˆ 3.94, P ˆ 0.033) could be explained with signi®cantly greater
MEPs for `listening to fairy-tales', `reading aloud' and `reading
silently' as compared with `white noise' (all t11  2.87, P < 0.02).
`Listening to fairy-tales' produced greater facilitation than `listening to
sentences', but the difference was not signi®cant (P ˆ 0.13).
Discussion
The present study demonstrated that productive and receptive lin-
guistic tasks excite the motor cortices for both hands. This facilita-
tion, which is below the threshold for overt movement generation,
was revealed by transcranial magnetic stimulation of cortical hand
ß 2003 Federation of European Neuroscience Societies, European Journal of Neuroscience, 18, 704±708
Language and the motor system 707
muscle representation. The effect was absent during non-linguistic
tasks.
Although not universally accepted, the idea that articulate language
evolved from manual gestures has been proposed many times (see
Corballis, 2002 for a detailed review). It is assumed that language
developed in several stages from a mirror system for grasping to a
complex imitation system for grasping, then from a manual-based
communication system to a proto-speech and ®nally to today's speech
and language (Arbib, 2000).
A series of experimental data has lent support to the plausibility of
this hypothesis. In monkeys the ventral premotor cortex (area F5)
contains neurons that discharge both when the monkey performs a
speci®c hand action and when it observes an individual executing a
similar action, the so-called `mirror neurons' (Di Pellegrino et al.,
1992; Rizzolatti et al., 1996a). Cytoarchitectonic studies suggest that
area F5 in the monkey is a homologue of part of Broca's region (area
44) in the human brain (Galaburda & Pandya, 1982). Cells in this area
might assist in the comprehension of other primates' motor acts.
Evidence from electrophysiological data likewise suggests that hand
actions are represented in Broca's area (Fadiga et al., 1995; Strafella &
Paus, 2000; Brass et al., 2001). Human brain imaging studies fre-
quently show coactivation of Broca's area during observation of hand
action (Buccino et al., 2001; Rizzolatti et al., 1996b; Nishitani & Hari,
2000).
Our data add to the existing experimental evidence on the gestural
theory of language origin by demonstrating a pre-excitation of the
primary hand motor cortex through speech. This indicates a direct link
between the language and manual/facial action systems, which is far
more extensive than previously thought and which may still be
functionally relevant in modern man. The study cannot prove the link
between the language and motor system, nor does it allow any ®rm
conclusions about the mechanisms by which language has developed.
Even though we have shown that speech perception activates the hand
motor area, the connection between speech and gestures might have
developed later in evolution.
The effects of language perception on oropharyngeal and facial
motor cortex excitability have recently been examined by Fadiga et al.
(2002). The authors found tongue motor cortex excitability to be
increased while subjects listened to speech. Their ®ndings support
the notion that the neural systems involved in perception of speech are
intimately linked with those involved in the execution of speech
(Liberman & Mattingly, 1985).
Our ®ndings expand their results by demonstrating that perception
of speech not only activates muscles directly involved in speech but
activates the motor system, in our study the hand gesture system, more
generally.
A previous study demonstrated that speech production activates the
hand motor cortex (Tokimura et al., 1996). This increase in hand motor
cortex activation might have been unspeci®c because articulation
activates the masseter muscle (Boroojerdi et al., 2000). The unique
®nding of the present study is that not only speech production but also
speech perception activates the hand motor cortex. Hand motor cortex
activation by mere language perception, as demonstrated in our study,
cannot easily be explained with an unspeci®c effect of subvocal
rehearsal particularly because other cognitively demanding auditory
and visuospatial tasks did not lead to a similar activation. A further
important difference to the study of Tokimura et al. (1996) is that we
found the hand motor cortex activation to be bilateral, contrary to their
lateralized MEP facilitation during reading and speaking aloud.
Although, in experiment I, we carefully reproduced their experimental
design, there are some differences that we believe make the present
study stronger. We only recruited subjects with known hemispheric
708 A. FloÈel et al.
language dominance. In the study of Tokimura et al. (1996), language
dominance in left-handed subjects was assumed to be right hemi-
spheric and vice versa. The probability of left-handed subjects being
right-hemispheric language dominant is, however, small (Knecht et al.,
2000b).
Bilaterality of activation is also observed in the majority of
functional imaging studies on language perception, usually with a
preponderance of the left hemisphere. However, some individuals
show lateralization of brain activation to the right hemisphere and
others fairly even activation of both hemispheres (Knecht et al.,
2000a,b). These activation patterns suggest that the blueprint of the
neural language system is bilateral for human species.
Our subhuman primate relatives, e.g. great apes, are fully capable of
complex motor planning and gestural communication (Hopkins &
Leavens, 1998). While showing some morphological asymmetries,
they lack distinct functional hemispheric specialization (Falk et al.,
1990; Cantalupo & Hopkins, 2001). Therefore, bilateral motor system
activation by phonetic gestures, as observed in our study, may repre-
sent a phylogenetically old mechanism that we still share with other
primates. It remains to be established whether there is already later-
alization at this level of development as suggested by the ®ndings by
Tokimura et al. (1996). We found no positive evidence but would not
rule out that other aspects of language than the ones tested here could
lead to lateralized activation of the hand motor system.
Our data demonstrate that speech leads to an automatic activation
not only of the oro-facial `gesture' systems (see Fadiga et al., 2002) but
also of the motor representations involved in gestures (hand motor
cortex). Moreover, this automatic activation takes place not only
during speech production (Tokimura et al., 1996) but also during
speech perception, which clearly does not involve manual gestures.
This language-to-motor excitation indicates that language is not
implemented in the brain as a separated module but is part of a
domain-general system (Hauser et al., 2002). Our ®ndings lend support
to the hypothesis that language may have evolved within an action-
perception network and possibly from a gestural communication
system (Corballis, 1992).
Acknowledgements
This work was supported by the Nachwuchsgruppen-FoÈrderung of the Ministry
of Science, Nordrhein-Westfalen (516-400 01000) and the Bennigsen-FoÈrder-
Preis of Nordrhein-Westfalen (IVA 6-400 302 97), the Innovative Medizinische
Forschung of the Medical Faculty of MuÈnster (Kn-1-1-II/96-34 and
KN 3 2 98 01) and the Deutsche Forschungsgemeinschaft, Bonn (Kn 285/4-1,
Kn 285/6-1 and Fl 379/1-1). We thank Sandra Kamping for her support in MEP
data analysis.
Abbreviations
LH-LD, left-handed left-hemispheric language dominant; LH-RD, left-handed
right-hemispheric language dominant; MEP, motor-evoked potential; RH-LD,
right-handed left-hemispheric language dominant; RH-RD, right-handed right-
hemispheric language dominant.
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