You are on page 1of 8

Neuropsychologia 40 (2002) 1379–1386

Laterality effects on motor awareness


Elena Daprati∗ , Angela Sirigu
Institut des Sciences Cognitives, 67 Bd Pinel, 69675 Bron, France
Received 26 January 2000; accepted 28 September 2001

Abstract
The ability to perceive self-produced movements and to correctly attribute an action to its proper agent is a natural task and a basic
requirement to human social communication. Recent experiments suggest that this apparently simple phenomenon is related to the mech-
anisms of motor production, raising the question whether recognition of self-produced movement is affected by asymmetries similar to
those present in motor skills. In this study, right- and left-handed subjects decided whether a moving hand presented on a screen was the
image of their own hand or of that of another person. Two experimental conditions were tested: either subjects saw their own moving
hand (subject condition) or they were shown the experimenter’s hand pantomiming their movement (experimenter condition); a glove
masked morphological differences between the two hands. Verbal responses and response times were analysed. Results showed that all
subjects were more accurate in recognising their preferred hand with respect to their non-preferred hand. Right-handers responded faster
than left-handers did, the latter group being especially slowed down in the experimenter condition. On the contrary, in the right-handers
group, response times did not differ among conditions. The present data indicate that the ability to recognise self-generated movements
is affected by motor dominance, thus suggesting that conscious knowledge of self-produced movements is closely related to the motor
system. © 2002 Elsevier Science Ltd. All rights reserved.
Keywords: Motor consciousness; Handedness; Motor skills; Manual asymmetry

1. Introduction An alternative interpretation suggests that hand preference


could be reflected by functional or structural asymmetries of
Over 90% of humans are right-handed, that is they show the primary motor cortex and descending pathways, either
a clear preference for the use of the right hand in the in isolation or in association with those suggested for higher
execution of skilled movements. Although this notion is order areas. In right-handers, a comparison of the relative
well-established and seems to be rather consistent across degree of activation between the two hemispheres following
cultures and over time [3,11], the neurobiological correlate ipsilateral movements reveals greater activation in the left
of handedness is still a matter of debate. Neuropsychological than in the right hemisphere [19], specifically in the region
evidence suggests that this behavioural lateralisation can be of the motor cortex [28]. Interestingly, this pattern seems
related to an asymmetry of the cortical areas controlling the to be reversed in left-handers. In a PET study, Kawashima
cognitive–motor requirements of skilled movements [14]. et al. [18] found that the right premotor cortex was acti-
As a general rule, movement of one limb is controlled by vated by both contralateral and ipsilateral finger movements,
the motor cortex of the contralateral hemisphere: however, whereas left premotor cortex, as well as primary motor cor-
at least in right-handers, a dominant role of the left hemi- tex and supplementary motor areas, were activated only by
sphere for praxis functions has frequently been invoked. contralateral movements.
For instance, ipsilateral deficits in tasks involving complex In a recent fMRI study, Dassonville et al. [6] have shown
cognitive–motor abilities have been described only follow- that the volume of activation in contralateral motor cortex
ing left (but not right) hemisphere damage [14]. Moreover, is greater during movements of the preferred hand in com-
ideomotor apraxia—i.e. a specific impairment in the per- parison to movements of the non-preferred hand in both
formance of complex gestures in the absence of elementary right- and left-handed subjects. A similar observation
sensory/motor deficits—is known to occur more frequently was obtained using whole-head magnetoencephalography
after left than after right hemisphere damage [12]. (MEG): Volkmann et al. [30] measured in vivo the hand rep-
resentation area in right- and left-handed subjects and found
∗Corresponding author. Tel.: +33-437911227; fax: +33-437911210. a significant expansion of hand motor cortex in the hemi-
E-mail address: daprati@isc.cnrs.fr (E. Daprati). sphere corresponding to the preferred hand. The authors

0028-3932/02/$ – see front matter © 2002 Elsevier Science Ltd. All rights reserved.
PII: S 0 0 2 8 - 3 9 3 2 ( 0 1 ) 0 0 2 0 3 - 2
1380 E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386

suggested that such an enlargement could provide the neural has been first described in the monkey’s ventral area 6, where
substrate for the more efficient processing of motor output neurones responding to both execution and observation of a
of the preferred hand. A recent study using transcranial mag- given action have been found [24]. In humans, although it
netic stimulation (TMS) supports this hypothesis [29]: mea- has been shown that the two systems, motor execution and
surable differences related to handedness have been found in motor representation, only partially overlap [10], the fact that
the excitability of the motor system. In particular, the thresh- similar cortical areas are activated while representing one’s
old for eliciting motor evoked potentials (MEPs) varies own action and observing someone else’s action implies that
according to hand dominance: lower TMS thresholds are re- the same representation may be shared among individuals.
quired for eliciting MEPs in the hemisphere contralateral to Self-consciousness might rely on the possibility to discrim-
the preferred hand. In other words, lower TMS thresholds for inate between central representations activated from within
one hand are associated with greater dexterity of that hand. (self-generated movement) from central representations ac-
Although the matter of motor asymmetry and hemispheric tivated from external agents (movement observation).
lateralisation has been extensively debated, few studies have Such a close link between execution and mental rep-
investigated this problem at the level of internal motor repre- resentation of movement raises the question whether the
sentations. A recent paper provided some evidence that the perception of self-performed actions can be influenced by
principle of contralateral control holds also for mentally sim- motor asymmetries. Recent experiments suggest that this
ulated movements [23]. Split-brain patients were required apparently simple phenomenon relies on a complex mech-
to judge handedness of line drawings depicting either right anism, which requires an efficient on-line control of motor
or left hands. Accuracy in the responses was higher when output [27] as well as the ability to update the body schema
the hand presented was contralateral to the perceiving hemi- in response to limb movements [5]. To our knowledge, no
sphere (e.g. a right hand presented to the isolated left hemi- study has yet addressed the question whether the capacity to
sphere), whereas patients performed at chance level when the recognise self-produced movements is subject to the same
stimulus was ipsilateral to the perceiving hemisphere. Data asymmetries present in motor skills.
collected in healthy subjects [16] further confirmed that an Sirigu et al. [27] required apraxic patients suffering from
advantage exists for each hemisphere when mental simula- left parietal lesion and age-matched normal controls to
tion involves movements of the contralateral hand compared identify the origin of a moving hand in an ambiguous situ-
to the ipsilateral one. Normal subjects required less time to ation. Both the preferred and the non-preferred hand were
select whether an overhand or an underhand grip was more tested. The results suggested that the ability to correctly
appropriate in order to grasp a manipulandum when the latter attribute a movement to its proper agent is reduced when
was presented to the hemisphere contralateral to the desig- subjects are asked to recognise the non-preferred hand. In
nated response hand. When a similar paradigm was admin- the present study, we further explore this issue in the at-
istered to a right-handed commissurotomised patient [17], a tempt to assess whether the ability to recognise one’s own
dissociation between the two hemispheres emerged. The left movement is subject to asymmetries related to motor dex-
(motor dominant) hemisphere was successful in represent- terity. Toward this end, both right- and left-handed subjects
ing with a high degree of accuracy both distal movements are required to execute hand gestures with their preferred
of the contralateral hand (i.e. grasping) and proximal move- and non-preferred hand and to monitor their execution on
ments of the ipsilateral hand (i.e. reaching). On the contrary, a TV screen. According to the experimental condition, the
the right hemisphere only appeared capable of representing image of the subjects’ hand can be substituted by that of the
distal movements of the contralateral hand with modest examiner’s, thus creating an ambiguous environment. The
accuracy, suggesting that processes needed to select upper subjects’ goal is to decide whether the presented hand be-
limb movements may be lateralised to the left hemisphere. longs to themselves or not. Indeed, if the ability to recognise
The presence of motor asymmetries in both real and men- one’s own movement is subject to the same asymmetries
tally simulated movements emerges even more directly in present in motor skills, recognition of the non-preferred
a recent study by Maruff et al. [20]. Fifty subjects (includ- hand should lead to an increased number of errors and/or
ing right- and left-handers) were required to either execute to longer decision time in both right- and left-handers.
or mentally simulate a visually guided pointing task [26].
Results showed a significant speed reduction when subjects
either used or imagined to use their non-dominant hand, thus 2. Methods
supporting the hypothesis that real and imagined movements
are controlled by similar constraints. However, asymmetries 2.1. Subjects
in both speed and error rate were greater for imagined move-
ments as compared to executed movements. This latter result Eighteen subjects participated to the experiment: nine of
is consistent with recent neuroimaging [13,15,25] and neuro- them (three men and six women, mean age 23.1 years, range
physiological [24] data suggesting that the link between exe- 21–27 years) were right-handers and nine were left-handers
cution and mental representation of movement is very tight, (two men and seven women, mean age 23.1 years, range
also extending to movement observation. A “mirror” system 18–38 years). Mean laterality coefficient, as assessed by the
E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386 1381

Fig. 1. Schematic representation of the apparatus: details in the text.

Edinburgh Inventory [22], was +0.87 for the right-handers (laterality coefficient 1.00), the other was almost ambidex-
and −0.75 for the left-handers. None of the subjects suffered trous (laterality coefficient 0.30).
from neurological or psychiatric pathologies. Vision was
normal or corrected to normal. Subjects were naive as to the 2.3. Procedure
purpose of the experiment.
Before the experimental session, subjects were shown the
2.2. Apparatus gestures they were to perform. Subjects started from the
fist-closed posture and moved their hand on the horizontal
A detailed description of the apparatus is given elsewhere plane, palm facing the table. Gestures required extension of
[4]. one or more digits, namely (1) thumb, (2) index, (3) thumb
Fig. 1 shows a schematic representation of the system: and index, (4) all fingers. The task for the subjects was to
subjects sat in front of a table on which a mirror was placed produce one gesture in response to a verbal command and to
(M), at about 35 cm from the subject’s frontal plane and monitor its execution on the screen in front of them. Subjects
inclined by 30◦ on the vertical plane. Subjects positioned knew in advance that in half of the trials the hand displayed
their hand below the mirror, on a comfortable support. A would be their own one (subject condition), whereas in the
camera (C1) filmed the subjects’ hand and a closed circuit other half, it would be that of another person (experimenter
television system reproduced it in order to make it visible to condition). In the latter condition, the examiner’s hand al-
the subjects on the mirror surface. Since no time delay was ways performed the same movement that was required to the
present, when the hand was placed below the mirror the im- subject. Subjects knew in advance that the two conditions
pression was that of seeing it through a window glass. An would be presented in random order. Indeed, the subjects’
identical set-up was located nearby and collected images of goal was to recognise whether the hand on the screen was
the experimenter’s hand (C2). During the experiment, either their own or not.
the image of the subject’s hand or that of the experimenter’s More details on the procedure are given in Fig. 2.
could be presented to the subject. Identical white gloves cov- Twenty-four trials were run for both the subject condition
ered both hands, in order to eliminate morphological differ- and the experimenter condition (six trials for each gesture
ences. In order to get a correct spatial and temporal match type). For each subject, both the right and the left hand were
between the subject’s and the experimenter’s hand, three tested in separate blocks: order of hand use was counterbal-
TV screens were available in the experimenter’s cabin. The anced among subjects.
monitors presented the images of both the subject’s and the Verbal responses were recorded: the trial was considered
experimenter’s hand, either individually or superimposed. successful when subjects responded “yes” to the image
In order to synchronise movement onset, both subject and of their own hand and “no” to that of the experimenter’s
experimenter started to move in response to one acoustic hand. A voice key system recorded response times. Re-
signal. All detectable differences in movement onset were sponse times shorter than 100 ms or longer than 4000 ms
controlled and trials in which timing or spatial matching was were considered errors and the corresponding trials were
imprecise were repeated at the end of the session. repeated at the end of the session. Due to the dichotomous
Two experimenters were sitting in the cabin: alternately, nature of the analysed variable, the number of correct re-
one acted as mime, while the other monitored task per- sponses was submitted to non-parameteric tests: namely
formance. One experimenter was strongly right-handed Mann–Whitney U-test for between groups comparisons and
1382 E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386

Fig. 2. Experimental procedure. From top to bottom: time = time-scale, screen = events appearing on the screen located in front of the subject,
task = instructions given to subjects. “Instruction” refers to the verbal command specifying the type of gesture subjects were required to perform.
“Movement” refers to the motor phase, elicited by an acoustic signal (beep). Subjects were allowed an interval of 2 s in order to perform the gesture
before vision of the hand was prevented. “Response” refers to the period of 4 s in which subjects were expected to give their answer.

Wilcoxon matched-pairs-tests for within groups compar- P = 0.004). This was true for both the condition in which
isons. Response times were analysed by means of three-way subjects saw their own hand (subject condition: T = 10.5,
ANOVA, group (right-handers versus left-handers), man- Z = 2.4, P = 0.01) and for the condition in which they saw
ual preference (preferred versus non-preferred hand) and the experimenter’s hand (experimenter condition: T = 27.5,
experimental condition (subject’s versus experimenter’s Z = 2.3, P = 0.02, see Table 1 for numerical values).
hand) being the main factors. Newman–Keuls-test was All subjects produced more correct responses in the sub-
used for post-hoc analysis on response times. Whenever ject condition, i.e. when they saw their own hand (subject
required, a Bonferroni correction for multiple comparisons condition 45.0, S.D. 2.2, experimenter condition 29.7, S.D.
was applied. 7.7; T = 0.0000001, Z = 3.62, P < 0.0003). In other
words, errors increased whenever ambiguity was introduced.
A closer look to the number of correct responses given
3. Results by each group of subjects showed that the advantage
for the preferred hand was significant only within the
All subjects well understood the task and errors in the right-handers group (correct responses: preferred 39.2, S.D.
procedure, i.e. execution of a movement different from that 3.3, non-preferred 36.7, S.D. 3.6; T = 0.00001, Z = 2.665,
requested, anticipation or delay in starting the movement or P < 0.01). The difference was not significant in the case of
giving the response were extremely rare (0.31% of trials) left-handers (preferred 37.8, S.D. 4.5, non-preferred 35.7,
and were randomly distributed among conditions. S.D. 4.5; P = 0.1, Fig. 3 left panel). A direct comparison
between the two groups failed to reach significance. As
3.1. Correct responses previously stated, all subjects produced more correct re-
sponses when they saw their own hand compared to when
Independently from their handedness, the 18 subjects they saw the experimenter’s hand. The decrease in the
produced significantly more correct responses when asked number of correct responses in the latter condition was sig-
to recognise their preferred hand with respect to the nificant for both right-handed subjects (experimenter con-
non-preferred hand (correct responses: preferred 38.5, S.D. dition 31.0, S.D. 7.4, subject condition 44.9, S.D. 2.3; T =
3.9, non-preferred 36.2, S.D. 4.0; T = 20.0, Z = 2.85, 0.00001, Z = 2.520, P < 0.01) and left-handed subjects

Table 1
Mean number of correct responses (S.D.) and mean response times (S.D.) for right- and left-handed subjects, and for the entire group
Preferred hand Non-Preferred hand

Subject’s hand Experimenter’s hand Subject’s hand Experimenter’s hand

Mean number of correct responses (S.D.) (maximum of correct responses = 24)


Right-handers 23.2 (0.8) 16.0 (3.7) 21.7 (1.6) 15.0 (3.8)
Left-handers 22.9 (0.9) 14.9 (4.4) 22.2 (1.9) 13.4 (5.0)
All subjects 23.0 (0.9) 15.4 (4.0) 21.9 (1.7) 14.2 (4.4)
Mean response times (S.D.) in ms
Right-handers 630.4 (153.1) 609.0 (160.1) 607.4 (211.9) 608.2 (183.5)
Left-handers 676.5 (264.1) 742.1 (301.7) 747.5 (192.8) 802.9 (224.9)
All subjects 653.4 (210.7) 675.5 (244.1) 677.5 (209.4) 705.5 (222.9)
E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386 1383

Fig. 3. Effect of manual preference on number of correct responses (left panel) and on response times (right panel) given by each group of subjects.
Preferred hand (black bars) refers to the right hand for right-handed subjects and to the left hand for left-handed subjects. The opposite for non-preferred
hand (dotted bars). Error bars indicate standard errors (preferred hand: continuous line, non-preferred hand: dotted line).

(experimenter condition 28.3, S.D. 8.3, subject condition (preferred versus non-preferred hand), experimental condi-
45.1, S.D. 2.2; T = 0.0001, Z = 2.665, P < 0.01). tion (subjects versus experimenter’s hand) and response ac-
curacy (correct versus incorrect answer) were within groups
3.2. Response times factors. No difference between the two groups was found. A
main effect of response accuracy emerged (F(1, 7) = 10.18,
When response times are considered, it clearly emerges P < 0.015), showing that average response times were
from Fig. 3 (right panel) that left-handed subjects were significantly slower when they led to incorrect (858.4 ms,
on average slower than right-handed subjects (left-handers S.D. 189.4) compared to correct responses (675.4 ms, S.D.
742.2 ms, S.D. 219.9; right-handers 613.8 ms, S.D. 141.7). 189.4). The interaction between experimental condition and
Although no main effect of group was found, a significant accuracy was also significant (F(1, 7) = 5.71, P < 0.0482):
interaction emerged between group and experimental con- namely, response latency significantly increased (926.3 ms,
dition (F(1, 16) = 4.515, P < 0.0495). Post-hoc analysis S.D. 258.4) when subjects saw their own hand but were
showed that left-handers were slower than right-handers in mistaken in recognising it (i.e. they responded “no”).
both the subject and the experimenter conditions (subject Summing up, the main results of the present study were
condition, right-handers: 618.9 ms, S.D. 150.4; left-handers: the following: (1) independently from their handedness, all
712.0 ms, S.D. 198.7, P < 0.001; experimenter condition, subjects provided more correct responses when they were
right-handers: 608.6 ms, S.D. 135.4; left-handers: 772.5 ms, asked to recognise their preferred hand. (2) All subjects
S.D. 247.9, P < 0.0002). Moreover, a consistent time in- produced more correct responses when they saw their own
crease between the “easier” subject condition (712.0 ms, hand compared to when they saw the experimenter’s hand,
S.D. 198.7) and the more ambiguous experimenter condition suggesting that errors increased when ambiguity was intro-
(772.5 ms, S.D. 247.9) was present only in the left-handers duced. (3) Left-handers were on average slower in providing
group. No difference in mean response times was found their response than were right-handed subjects. (4) Longer
among right-handers (Fig. 3 right panel, Table 1). response latencies were not associated to correct responses.
Further analyses were run in order to test any evidence
for a speed–accuracy trade-off in subjects’ performance. 4. Discussion
Response times for correct responses were compared to re-
sponse times for incorrect responses by means of a four-way Most people have a preferred hand, whose skilfulness
ANOVA. Group (right-handers versus left-handers) was largely exceeds that of the so-called non-preferred hand.
the between groups factor whereas manual preference Asymmetries related to motor dominance have been de-
1384 E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386

scribed to affect imagined movements [20]. The present least two non-conflicting interpretations. Either subjects are
study was designed in order to assess whether this notion ex- more trained to attend to their preferred hand during praxis,
tends to the capacity to recognise self-produced movements, or self-awareness of motor outputs is richer for the more
or in broader terms, to self-awareness in motor acts. Right skilled hand. According to the first interpretation, we can
and left-handed healthy subjects were required to produce a speculate that the better performance for the dominant hand
hand gesture in an ambiguous environment, in which direct could be a secondary effect of familiarity, subjects having
vision of their moving limb was prevented and visual feed- acquired major expertise in visually controlling movements
back provided on a TV screen could be manipulated. Sub- of their preferred hand. On the other hand, it could not be
jects were asked to decide whether they were the agents of excluded that, when attempting to recognise their own hand,
the presented movements. Results showed that the number subjects made use of an internal representation of their dom-
of correct responses given by the subjects varied according inant hand that is “motor” in nature. Evidence has been
to the hand used. Both right- and left-handers were more provided that motor habits can influence our perception of
accurate when asked to identify their preferred hand with the visual world. Objects that can be grasped and manipu-
respect to their non-preferred hand, suggesting the presence lated appear to be processed also in relation to the manual
of a close link between motor dominance and motor aware- dominance of the subject, thus suggesting the involvement
ness. We will define here motor awareness as the ability of of motor processes (namely motor imagery) in non-motor
an individual to both perceive and visually recognise a given functions [7]. In the present task, subjects could have found
movement as his/her own. In other words, as the knowledge recognition of the preferred hand easier either due to a more
enabling subjects to state that they are the source and the familiar visual image or to a richer visual–motor knowledge
owner of a presented gesture. This ability necessarily comes of their more skilled hand. We favour this second hypothesis,
into play in the task used in the present experiment. which points to a close relation between the system respon-
As previously reported [27], the accomplishment of the sible for motor consciousness and the structures involved in
present task likely requires several steps. First, in addition motor representation and movement production.
to intentional information, subjects must be able to form What kinds of mechanisms allow the interplay between
a reliable representation of the movement they are about consciousness of movement and motor production? Manual
to execute. More precisely, subjects should be able to pre- preference has an anatomic and functional counterpart in
dict the proprioceptive and visual consequences of their the expansion of hand motor cortex for the preferred hand
act and to fit actual incoming kinaesthetic information into [1,30] and this phenomenon has been invoked to explain the
their predictive model. As a result, they will “know” which more efficient processing of the corresponding motor out-
movement they are executing and whether it matches their put. Efficiency could improve thanks to an increase in the
intentions. Secondly, this information is compared to the number of neural elements and/or tighter connectivity: mon-
one provided by the visual system, thus making possible to key data show both an enlargement and an increased degree
decide whether the movement shown to the subjects belongs of spatial complexity of motor areas opposite to the pre-
to themselves or not. Once the result of this comparison is ferred hand [21]. At the cortical level, this modification may
made conscious, a response can be produced. account for better interaction between neuronal assemblies
We recently provided some evidence that brain damage representing muscles that are used in close spatial or tem-
can disrupt this motor recognition mechanism at the level of poral contiguity. Analysis of cytoarchitectonic differences
action representation, thus preventing subjects from having between the two hemispheres [1] have shown an increased
a reliable image of the movement they are executing. More neuropil compartment in BA 4 in the dominant hemisphere,
particularly, in apraxic patients suffering from left parietal supporting the hypothesis that hand preference is associated
lesion [27], the ability to either form or update a reliable with increased connectivity. At the behavioural level, such
predictive model of the executed gestures is impaired. Ac- improved interaction may be responsible for a more skilled
cordingly, when attempting to reproduce a complex hand motor repertoire of the preferred hand. In agreement with
posture, these patients fail in their execution as well as in the above data, the present results further indicate that not
recognising their own movement. only motor production but also motor awareness can ben-
The present results on healthy subjects support the hy- efit from the anatomical and physiological pattern linked
pothesis that the ability to visually recognise a movement as to hand preference. Handedness asymmetries seem, thus,
one’s own is subject to asymmetries similar to those present to extend beyond the level of primary motor functions and
in motor skills. It is unlikely that our results may be ac- likely emerge also in those structures which are involved in
counted for by the possible time delay intervening between cognitive–motor operations.
movement executed and movement observed by the sub- Two more points emerged from the results of the present
jects. Indeed, had this been the cue to the correct answer, it experiment. First, the difference between preferred and
should have produced an equal number of correct responses non-preferred hand was less evident in the left-handers
for the preferred and non-preferred hand. group. Second, left-handers were on average slower in
The increase in the number of correct responses after pre- giving their responses than right-handers. The former re-
sentation of the dominant hand could be accounted for by at sult could be accounted for by the fact that left-handers
E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386 1385

are more heterogeneous and less markedly lateralised than References


right-handers: for instance, on preference measures, i.e.
hand or foot usage, left-handers show larger variability [1] Amunts K, Schlaug G, Steinmetz H, Dabringhaus A, Roland PE,
and lower lateralisation than right-handed subjects [2]. In Zilles K. Asymmetry in the human motor cortex and handedness.
Neuroimage 1996;4:216–22.
addition, in hand selection responses to unimanual tasks,
[2] Borod JC, Caron HS, Koff E. Left-handers and right-handers
left-handers do not respond with the preferred hand as con- compared on performance and preference measures of lateral
sistently as do right-handers [8]. This reduced lateralisation dominance. British Journal of Psychology 1984;75:177–86.
emerged also in the sample examined in the present experi- [3] Coren S, Porac C. Fifty centuries of right-handedness: the historical
ment: only one of our left-handed subjects scored between record. Science 1977;198:631–2.
−0.90 and −1.00 at the Edinburgh Inventory, whereas five [4] Daprati E, Franck N, Georgieff N, Proust J, Pacherie E, Dalery J,
et al. Looking for the agent. An investigation into consciousness of
out of nine right-handed subjects showed such an extreme action and self-consciousness in schizophrenic patients. Cognition
score. Indeed the absolute scores at the handedness test 1997;65:71–86.
differed significantly between the two groups (U = 18.00, [5] Daprati E, Sirigu A, Pradat-Dihel P, Franck N, Jeannerod M.
Z = 1.987, P = 0.046) Accordingly, we can assume that Recognition of self-produced movement in a case of severe neglect.
in our sample the reduced asymmetry in hand motor skills Neurocase 2000;6:477–86.
[6] Dassonville P, Zhu XH, Ugurbil K, Kim SG, Ashe J. Functional
present in the left-handers group could be responsible for activation in motor cortex reflects the direction and the degree of
the reduced handedness effect. Both results are consistent handedness. Proceedings of the National Academy of the Sciences
with a previous report by Gentilucci et al. [9] in which USA 1997;94:14015–8.
right- and left-handed subjects were required to recog- [7] de’Sperati C, Stucchi N. Recognizing the motion of a graspable
nise whether a static hand holding an object, with either object is guided by handedness. Neuroreport 1997;8:2761–5.
[8] Gabbard C, Iteya M, Rabb C. A lateralized comparison of handedness
a congruent or incongruent type of grip, corresponded to
and object proximity. Canadian Journal of Experimental Psychology
the picture of a right or a left hand. Right-handers were 1997;51:176–80.
faster in judging right hands, whereas no difference was [9] Gentilucci M, Daprati E, Gangitano M. Right-handers and
found between preferred and non-preferred hand in the left-handers have different representations of their own hand.
left-handed subjects. Moreover, left-handers were on av- Cognitive Brain Research 1998;6:185–92.
erage slower than right-handers. A similar pattern is also [10] Gerardin E, Sirigu A, Lehericy S, Poline JB, Gaymard B, Marsault
C, et al. Partially overlapping neural networks for real and imagined
found in another previous report. When asked to decide hand movements. Cerebral Cortex 2000;10:1093–4.
whether a rotating screwdriver was either screwing or un- [11] Gilbert AN, Wysocki CS. Hand preference and age in the United
screwing [7], healthy subjects required longer response States. Neuropsychologia 1992;30:601–8.
times in those conditions implying motor imagery of the [12] Gonzales-Rothi LJ, Heilman KM, editors. Apraxia: the
non-dominant hand. Moreover, left-handers were overall neuropsychology of action. New York: Lawrence Erlbaum,
1996.
slower than right-handers. As suggested by Gentilucci et al.
[13] Grezes J, Costes N, Decety J. The effects of learning and intention
[9], the behaviour of left-handers could be explained by the on the neural network involved in the perception of meaningless
fact that, when required to covertly represent movement actions. Brain 1999;122:1875–87.
of their own hands, they likely rely on a “pictorial” (i.e. [14] Haaland KY, Harrington DL, Clinical implications of ipsilateral
more observation-based) rather than “pragmatic” (i.e. motor motor deficits after unilateral hemispheric damage. In: Bruno A,
Chollet F, Vellas BJ, Albarede JL, editors. Facts and research
experience-based) image of their dominant hand. In agree-
in gerontology: stroke in the elderly. New York: Springer, 1996:
ment with this hypothesis, we can assume here that, on pp. 101–14.
the one hand, left-handers are less affected by asymmetries [15] Iacoboni M, Woods RP, Brass M, Bekkering H, Mazziotta JC,
related to motor dexterity. On the other hand, they likely Rizzolatti G. Cortical mechanisms of human imitation. Science
need further steps in the analysis of the presented image, 1999;286:2526–8.
such as a transition between a visual and a motor-based [16] Johnson SH. Cerebral organization of motor imagery: Contralateral
control of grip selection in mentally represented prehension.
representation, in order to access those subtle kinematics Psychological Science 1998;9:219–22.
cues which are required to recognise one’s own movement. [17] Johnson SH, Corballis PM, Gazzaniga MS. Within grasp but out of
In conclusion, we think that the present data demonstrate reach: Evidence for a double dissociation between imagined hand and
how the ability to recognise self-generated movements can arm movements in the left cerebral hemisphere. Neuropsychologia
be affected by motor dominance, suggesting that the system 2001;39:36–50.
[18] Kawashima R, Inoue K, Sato K, Fukuda H. Functional asymmetry
responsible for motor awareness is profoundly nested within
of cortical motor control in left-handed subjects. Neuroreport
the mechanisms controlling motor production. 1997;8:1729–32.
[19] Kim SG, Ashe J, Hendrich K, Ellerman JM, Merkle H, Ugurbil
K, et al. Functional magnetic resonance imaging of motor
Acknowledgements cortex: hemispheric asymmetry and handedness. Science 1993;261:
615–7.
[20] Maruff P, Wilson PH, De Fazio J, Cerritelli B, Hedt A, Currie J.
The authors wish to thank Virginie Czernecki for her pre- Asymmetries between dominant and non-dominant hands in real and
cious help in collecting the data. This work was supported imagined motor task performance. Neuropsychologia 1999;37:379–
by CNRS. 84.
1386 E. Daprati, A. Sirigu / Neuropsychologia 40 (2002) 1379–1386

[21] Nudo RJ, Jenkins WM, Merzenich MM, Prejean T, Grenda [26] Sirigu A, Duhamel JR, Cohen L, Pillon B, Dubois B, Agid Y.
R. Neurophysiological correlates of hand preference in primary The mental representation of hand movements after parietal cortex
motor cortex of adult squirrel monkeys. Journal of Neuroscience damage. Science 1996;273:1564–7.
1992;12:2918–47. [27] Sirigu A, Daprati E, Pradat-Dihel P, Franck N, Jeannerod M.
[22] Oldfield RC. The assessment and analysis of handedness: the Perception of self-generated movement following left parietal lesion.
Edinburgh inventory. Neuropsychologia 1971;9:97–113. Brain 1999;122:1867–74.
[23] Parsons LM, Gabrieli JD, Phelps EA, Gazzaniga MS. Cerebrally [28] Taniguchi M, Yoshimine T, Cheyne D, Kato A, Kihara T, Ninomiya
lateralized mental representations of hand shape and movement. H, et al. Neuromagnetic fields preceding unilateral movements in
Journal of Neuroscience 1998;18:6539–48. dextrals and sinistrals. Neuroreport 1998;9:1497–502.
[24] Rizzolatti G, Fadiga L, Gallese V, Fogassi L. Premotor cortex and the [29] Triggs WJ, Calvanio R, Levine M. Transcranial magnetic
recognition of motor actions. Cognitive Brain Research 1996;3:131– stimulation reveals a hemispheric asymmetry correlate of intermanual
41. differences in motor performance. Neuropsychologia 1997;35:
[25] Rizzolatti G, Fadiga L, Matelli M, Bettinardi V, Paulesu E, Perani 1355–63.
D, et al. Localization of grasp representations in humans by [30] Volkmann J, Schnitzler A, Witte OW, Freund HJ. Handedness and
PET: 1. Observation versus execution. Experimental Brain Research asymmetry of hand representation in human motor cortex. Journal
1996;111:246–52. of Neurophysiology 1998;79:2149–54.

You might also like