Neuropsychologia 43 (2005) 227–237

Motor awareness without perceptual awareness!

Helen Johnson, Patrick Haggard∗
Institute of Cognitive Neuroscience and Department of Psychology, University College London,
Alexandra House, 17 Queen Square, London WC1N 3AR, UK

Abstract

The control of action has traditionally been described as “automatic”. In particular, movement control may occur without conscious
awareness, in contrast to normal visual perception. Studies on rapid visuomotor adjustment of reaching movements following a target shift have
played a large part in introducing such distinctions. We suggest that previous studies of the relation between motor performance and perceptual
awareness have confounded two separate dissociations. These are: (a) the distinction between motoric and perceptual representations, and
(b) an orthogonal distinction between conscious and unconscious processes. To articulate these differences more clearly, we propose a new
measure of motor awareness, based on subjects’ ability to reproduce the spatial details of reaching movements they have just made. Here
we focus on the dissociation between motor awareness and perceptual awareness that may occur when subjects make rapid visuomotor
adjustments to reaching movements following a target shift. In experiment 1, motor awareness was dissociated from perceptual awareness of a
target shift during reaching movement. Participants’ reproduction of movement endpoints following visuomotor adjustment was independent
of whether they saw the target shift or not. Experiment 2 replicated this result, and further showed that neither motor awareness nor motor
performance were disrupted by TMS over the parietal cortex. The neural mechanisms underlying motor awareness, and the implications for
theories of consciousness, are discussed.
© 2004 Elsevier Ltd. All rights reserved.

Keywords: Human; Reaching movement; Unconscious processing; Consciousness; Transcranial magnetic stimulation

1. Introduction work was continued over several years by Prablanc, Jean-

Studies of the neural mechanisms underlying coordinated
action have traditionally occupied a curious middle ground
between psychology and neuroscience. This has given the
field a very productive, interdisciplinary flavour, and allowed
researchers to raise strictly scientific questions, which at the
same time seem of fundamental importance to the wider view
of the mind and human nature. This combination of scien-
tific knowledge and a willingness to tackle major intellectual
questions in a clear and practical way is very characteristic
of the work of Marc Jeannerod, throughout his distinguished
career.
In this paper, we particularly focus on the relation between
skilled action and conscious experience. Marc Jeannerod was
among the first to investigate this relation scientifically. The
! This study was carried out at the Institute of Cognitive Neuroscience.
∗ were able to correct their ongoing trajectories according to the
Corresponding author. Tel.: +44 20 7679 1151, fax: +44 20 7813 2835.
E-mail address: p.haggard@ucl.ac.uk (P. Haggard).
nerod and others, in an extensive series of studies using the
“double-step paradigm” of visuomotor adjustments in reach-
ing movements (e.g. Pélisson, Prablanc, Goodale, & Jean-
nerod, 1986; Prablanc & Martin, 1992). The double-step
paradigm was initially studied in saccadic eye movements
(Hallet & Lightstone, 1976) but we focus here on the clas-
sic result in the case of reaching movements. Pélisson et al.
(1986) asked subjects to make rapid pointing movements to-
wards a visual target, without vision of the hand. In their
paradigm, subjects initially fixated a peripheral light, and
made a saccade to the target when it first appeared, and then
reached towards it (step 1). During the saccade, the target was
unpredictably shifted a small distance either to the left or to
the right (step 2). Because of the known reduction in visual
perception during a saccade (Matin, 1974), subjects were not
perceptually aware that the target had shifted. Nevertheless,
inspection of the movement trajectories showed that subjects
direction of the visual target shift. That is, the brain systems

0028-3932/$ – see front matter © 2004 Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuropsychologia.2004.11.009
228 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237
generating motor awareness clearly had access to informa-
tion about the target location that did not enter into perceptual
awareness.
Since this early work, several additional studies by
Marc Jeannerod and others have replicated the basic result
(Castiello, Paulignan, & Jeannerod, 1991; Day & Brown,
2001; Day & Lyon, 2000; Desmurget et al., 1999; Paulignan,
MacKenzie, Marteniuk, & Jeannerod, 1991; Prablanc & Mar-
tin, 1992). This work became widely known and had strong
impact on our way of thinking about the mind and brain in
general. For example, the basic finding drove the develop-
ment of the distinctions between the dorsal and ventral visual
pathways (Goodale & Milner, 1992), and between the brain
systems for semantics and pragmatics (Jeannerod, 1994).
Many studies using the double-step paradigm have trig-
gered the target shift directly from the saccade, in order to
prevent perceptual awareness of the target shift. Relatively
few have measured perceptual awareness directly, and even
fewer have investigated what factors influence perceptual
awareness. Therefore, the initial claim of dissociation be-
tween motor performance and perceptual awareness was not
examined in detail. An important exception, however, was the
study by Castiello et al. (1991). They used the basic double-
step paradigm, as before. Unusually, they did not attempt
to prevent vision of the target shift. Instead, they required
subjects to respond to the target shift in different ways in
each of three conditions. The first condition replicated the
double-step paradigm: subjects responded to the target shift
by adjusting an ongoing reaching movement. In the second
condition, subjects did not move their hand, but made a sim-
ple vocal response on detecting the target shift. In the final
condition, subjects made both of these responses on each
trial. Castiello et al. argued that the simple verbal response
required perceptual awareness of the target shift, whereas a
visuomotor adjustment did not.
The results showed that visuomotor adjustments occurred
within approximately 100 ms of the target shift, while ver-
bal responses took over 300 ms. This latency difference was
present whether the responses were made on the same trial
(third condition) or individually (first and second conditions).
The authors interpreted the delay in the verbal response
as reflecting the slow operation of the brain circuits me-
diating perceptual awareness. The circuits underlying con-
scious processing are either additional to, or distinct from
and slower than, circuits underlying visuomotor responses.
This result provides convergent behavioural evidence for
a dissociation between motor performance and perceptual
awareness.
While such studies have been very influential, and have
shed new light on the brain’s visuomotor systems, they do not
conclusively reveal the relations between action and aware-
ness. These studies appear to dissociate processes, which in
fact differ in at least two respects. That is, motor performance
differs from perceptual awareness in that the former is mo-
tor and the latter perceptual, but also in that the former is a
performance or behaviour, while the latter is an awareness,
or subjective state. Dissociating processes which differ in
more than one respect is problematic: which dimension of
difference is responsible for the differences seen in experi-
mental data? Moreover, processes can only be dissociated if
they were previously accepted as unitary. As the number of
attributes along which the processes are accepted to differ
increases, an approach based on dissociation becomes less
convincing.
We therefore suggest that the scientific issues raised by the
double-step paradigm can be broken down into investigations
of two separate dissociations. First, does motor performance
dissociate from motor awareness? That is, what do people
consciously know about their own visuomotor adjustments?
Second, does motor awareness dissociate from perceptual
awareness? That is, can one be aware of a visuomotor ad-
justment even when unaware of any target shift? We recently
gave an affirmative answer to the first question (Johnson, van
Beers, & Haggard, 2002). By asking subjects to reproduce
the spatial path of a movement they had just made in the
double-step paradigm, we found that subjects typically un-
derestimated the latency and the gain of their own visuomotor
adjustments. In an anti-point condition, however, both these
results were reversed. This pattern of results suggested that
the awareness subjects had of their visuomotor adjustment
depended strongly on the role of intention in generating the
adjustment. Motor awareness followed motor intention. We
use the term “motor awareness” here to indicate subjects’
conscious representation of their own action, effectively as
an opposite to the traditional use of the term “automatic” in
the psychology of action.
Here, we consider the second of the two dissociations pro-
posed above: how is motor awareness related to perceptual
awareness? In the double-step reaching task, subjects may or
may not be aware of the target shift on any particular trial,
according to how successfully the paradigm implements sac-
cadic suppression. At the same time, subjects may have full,
limited, or no awareness that their movement has deviated
to one side (Johnson et al., 2002). In principle, these two
varieties of awareness should be interdependent because the
spatial path of a movement towards the target physically de-
pends on the spatial location of the target. For example, if the
subject sees the target jump to one side, and is trying to reach
for that target, they should correct their movement, and should
be aware that they have done so. Conversely, if the subject be-
comes aware that their movement trajectory has veered to one
side, it seems logical to conclude that the trajectory deviated
in pursuit of a target, which had shifted to that side. In general,
a consciously perceived target shift should be associated with
awareness of any resulting motor adjustment. However, the
psychological modules responsible for perceptual and motor
awareness may not follow the strict dependency of the spatial
world: indeed dissociative disorders are an established feature
of human consciousness (Farah & Feinberg, 2000). Motor
awareness and perceptual awareness could be a dissociable
attributes of the human mind, subserved by separate brain
structures.
 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237
Therefore, we performed two experiments to study the
relationship between perceptual awareness and motor aware-
ness in further detail. In experiment 1, we combined saccadic
suppression with a double-step reaching task. We asked sub-
jects to reproduce each movement as a measure of motor
awareness, and we considered how motor awareness var-
ied between trials where subjects had reported perceptual
awareness of the target shift and those where they had not.
In experiment 2, we attempted to disrupt visuomotor ad-
justments using TMS over the parietal cortex in a similar
paradigm.
2. Experiment 1
2.1. Methods
The details of the method and apparatus were based on pre-
vious studies (Desmurget et al., 1999; Pélisson et al., 1986).
Fifteen right-handed subjects (10 female, mean age 26 years)
participated with informed consent. Subjects sat at a table in
a darkened room with their head on a chin rest. The right hand
was occluded. Visual targets (7 mm diameter) were projected
onto a horizontal plane above the table, which the subject
viewed via a mirror, so that they appeared to be in front of
the hand. The central (unperturbed) target was aligned in the
parasagittal plane 14 cm in front of the start position. Two
other targets were at 8 cm to either side of the central target
in the frontoparallel plane. When the central target ‘jumped’
it moved 4 cm horizontally in either direction (see Fig. 1).
An initial visual fixation point was shown 40 cm to the left
of the central target. Reaching movements were recorded at
120 Hz with an electromagnetic tracker mounted on the index
fingernail. Movement start and end were defined as the point
at which the velocity of the finger fell below 5 cm/s.
Subjects performed 10 practice trials, then 240 experi-
mental trials in blocks of 20. In 60 trials, the central target
illuminated; in 50 trials, the left target illuminated; and in 50
trials, the right target illuminated. In a further 40 trials, the
centre target appeared and then jumped left, and in 40 trials,
it jumped right. Trial order was randomised across subjects.
Fig. 1. Experimental set-up (note: targets were 7 mm in diameter, but are shown enlarged here for clarity).
229
Each trial had three phases. In the first, reaching phase,
the subject initially fixated the peripheral spot for 2–3 s. The
fixation spot disappeared and one of the three targets im-
mediately appeared. Subjects looked at, and then reached
for the illuminated target as quickly as possible. During the
saccade period (200–300 ms after target onset) the central
target might jump unpredictably by 4 cm to the left or right
(Fig. 1). At the end of the movement, the target was extin-
guished and subjects returned to the starting position. This
initiated the second, reproduction phase of each trial. A 2 mm
wide guideline was shown across the entire table from left to
right, at the level of the targets. Subjects were asked to re-
peat the spatial detail of the movement path that they had
just made. They were specifically instructed to reproduce
the spatial features, without regard to movement speed or
other temporal information. During the movement reproduc-
tion phase, a guideline at the Y location of the targets was
projected to indicate the appropriate movement amplitude,
but no targets were visible. At the end of the reproduced
movement, the guideline disappeared and subjects again re-
turned to the starting point. Third and finally, subjects were
asked to verbally report any target jump (saying ‘left’, ‘right’
or ‘no’) seen during the reaching phase of the trial, as a mea-
sure of perceptual awareness. The full sequence of events is
shown in Fig. 2.
3. Experiment 1
3.1. Results
Subjects reported perceptual awareness of a target shift on
37% of jump trials.
Only trials in which the central target was initially illumi-
nated were analysed kinematically. Of these 6.34% of trials
were omitted due to equipment failure. A 2.11% of trials were
omitted due to pre-emptive eye or hand movements. A 7.8%
of trials were omitted as false positives (in non-jump trials)
or movement in the wrong direction (in jump trials). Exam-
ple traces for one participant are shown in Fig. 3. The initial
trial and the subsequent trajectory reproduction are shown
230 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237

Fig. 2. Events occurring in each trial.

for one trial on which the subject reported seeing the target
shift, and a second trial on which they did not. The y compo-
nent of the trajectories shows the smooth reaching movement
seen in earlier studies. The movement duration was typically
around 600 ms. The x component shows a clear movement ad-
justment towards the target, which occurs during the reach-
ing movement, and without pause or reprogramming. This
begins around 150 ms after movement onset. Note that the
adjustment is similar in trials with and without perceptual
awareness of the target shift.
We initially hoped to compare the full spatial paths of
initial and reproduced movements, in order to investigate
whether motor awareness of visuomotor adjustments. How-
ever, inspection of the results revealed a strong rightward bias
in the initial reaching movements. This was substantially re-
duced in the reproduced movements. The difference in bias
tended to obscure the small lateral deviations, which mark the
onset of the visuomotor adjustment. Therefore, we preferred
to study the final hand positions, by which point the visuo-
motor adjustment was easily detectable despite the bias. We
discuss this bias in more detail below.
Our main analysis therefore focussed on the final position
of the hand in the x direction. In the reaching phase of the
trial, this provides a measure of visuomotor adjustment fol-
lowing a target jump. Trials in which the target jumped were
divided into those in which the subject detected the jump and
those in which they did not (unaware), and these were anal-
ysed separately. Endpoints of all jump trials can be seen in
Table 1.
3.2. Endpoints of initial reaching movements
We first analysed the initial movements to detect whether
visuomotor adjustments to the target jump occurred. The
mean endpoint of each perturbed condition was compared
to the mean endpoint of the equivalent unperturbed condi-

Table 1
Mean and S.D. across subjects of endpoint × deviation for initial and reproduced movements (mm)
Endpoint deviations Left (target = −40 mm) Centre (target = 0 mm) Right (target = 40 mm)

Aware Unaware n/a Aware Unaware

Initial movement
Mean (mm) −9 −11 25 57 63
S.D. (mm) 29 30 30 36 34
Reproduced movement
Mean (mm) −28 −32 14 55 58
S.D. (mm) 32 29 29 31 33
Note: Negative numbers indication mm left of centre, positive numbers are mm right of centre.
 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237 231

Fig. 3. Typical perturbation (“jump”) trials for one participant showing the trajectories for their initial and subsequently reproduced (“copied”) movements.
The upper row shows a trial after which the subject reported seeing the target shift. In the trial in the lower row, the subject reported no change in target position.

tion. Family wise Bonferroni corrections were performed,
and so the significance level for the following t-tests is 0.025.
All movement endpoints differed significantly from the un-
perturbed trials regardless of whether the subject was aware
(left: t14 = 11.43, P > 0.001, right: t14 = −8.07, P > 0.001)
or unaware (left: t14 = 16.75, P > 0.001, right: t14 = −21.41,
P > 0.001) of the target shift. The effect of awareness on ad-
justment amplitude was then considered. No significant dif-
ference was found between the endpoints of aware and un-
aware trials for jumps to either the left (t14 = 1.54, P = 0.15)
or the right (t14 = −0.58, P = 0.57). That is, subjects were
just as efficient in adjusting their movements following tar-
get jumps regardless of whether they were aware of the jump
or not: motor performance was independent of perceptual
awareness. The average endpoints to targets that jumped to
the left were 10 mm to the left of centre. Average endpoints
to targets that jumped to the right were 60 mm to the right of
centre. Thus, there was an overall rightward bias (see above),
but also clear evidence of a normal visuomotor adjustment
pattern in the initial movements.
3.3. Endpoints of reproduced movements
We then investigated whether subjects were aware of their
visuomotor adjustments by repeating the same comparisons
for the reproduced movements. The reproduced endpoints of
each perturbed condition were compared to the reproduced
endpoints in the unperturbed condition. Following family
wise Bonferroni corrections, the significance level for the
following t-tests is again 0.025. All reproduced endpoints
on perturbed trials differed significantly from those on un-
perturbed trials regardless of whether the subject was aware
(left: t14 = 10.06, P > 0.001, right: t14 = −11.36, P > 0.001)
or unaware (left: t14 = 17.26, P > 0.001, right: t14 = −15.78,
P > 0.001) of the target shift. The effect of awareness on
the extent of the reproduced adjustments was then consid-
232 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237
ered. No significant difference was found between the repro-
duced endpoints of aware and unaware trials for jumps to
either the left (t14 = −1.20, P = 0.25) or the right (t14 = 1.28,
P = 0.22).
The relationship between initial endpoints and reproduced
movements in jump trials gives a measure of motor awareness
(see Table 1). Inspection of the table shows that the subjects
accurately reproduced the direction of their visuomotor ad-
justments in the immediately preceding movement.
To investigate the relation between perceptual awareness
and motor awareness, a 2 × 2 × 2 ANOVA was conducted on
the perturbation trial data (see Fig. 4). The factors were direc-
tion of target jump (left, right), perceptual awareness (aware,
unaware) and phase of trial (initial movement, reproduced
movement).
There was a significant effect of side (F1,14 = 265.96,
P > 0.001). On average, endpoints to leftward target shifts
were 20 mm left of centre, and 59 mm right of centre for
rightward shifts. There was no significant effect of aware-
ness (F1,14 = 0.35, P = 0.56) There was a significant effect of
trial phase (F1,14 = 22.49, P > 0.001). This occurred because
the rightward bias previously mentioned in the initial move-
ments was reduced in the reproduced movements. A signif-
icant interaction between side and movement (F1,14 = 24.1,
P > 0.001) indicated that this reduction was most marked for
leftward target jumps. There was no significant interaction
between trial phase and awareness (F1,14 = 2.92, P = 0.11),
nor awareness and side (F1,14 = 1.91, P = 0.19). Critically,
there was no interaction between side, awareness and trial
phase (F1,14 = 0.015, P = 0.91). That is, subjects’ ability to
reproduce the spatial detail of visuomotor adjustments was
equivalent in trials with and without perceptual awareness of
the target jump.
4. Experiment 1
4.1. Discussion
4.1.1. End points of initial movements
The mean endpoint of the movements during which the
target jumped were significantly different from the mean
Fig. 4. Mean endpoints in perturbation (“jump”) trials (mm).
endpoint of the central trials. That is, subjects performed
functional visuomotor adjustments following the target shift.
However, no significant difference was found between the
endpoints for those trials in which the subject was aware of
the target shift and those in which they were not. This con-
firms previous findings that efficient adjustments can be made
to an ongoing movement in response to a target shift regard-
less of whether the subject is aware of that target shift or not
(Day & Lyon, 2000; Desmurget et al., 1999; Paulignan et al.,
1991; Pélisson et al., 1986). The above studies have generally
studied manual responses to targets that shift during saccadic
suppression of conscious experience. Therefore, the dissoci-
ation between manual performance and visual awareness that
they report correlates with a difference between effectors: the
hand and arm in one case, and the oculomotor system in the
other. However, the dissociation between performance and
awareness can be found even in a single effector. Gaveau et al.
(2003) recently reported that subjects adjusted saccade kine-
matics for small changes in visual target position of which
they were perceptually unaware.
Returning to the present experiment, initial movements in
all conditions were subject to a rightward bias of approxi-
mately 2 cm. Such biases have also seen in other similar ex-
periments (Desmurget et al., 1999). We believe it may reflect
biomechanical factors (Carey et al., 1996; Carey & Otto de
Hart, 2001), coupled with the lack of visual feedback of hand
position. We speculate that the visual guide presented during
the reproduction phase of the movement may have helped to
reduce this bias.
4.1.2. Reproduced movements and motor awareness
The most important results concern motor awareness. Sub-
jects reproduced the spatial details of the visuomotor adjust-
ment on the immediately preceding trial. A similar finding
had been reported previously (Johnson et al., 2002). How-
ever, in that earlier study, subjects had full vision of the limb
throughout both the initial and the reproduced movement.
Therefore, the reproduced movements in that study could
reflect a visual memory of limb positions during the initial
movement, as well as, or instead of a representation truly
linked to movement. Moreover, that study focussed on the
perceived latency of visuomotor adjustments, whereas the
present analysis of endpoints focuses on perceptual gain. Our
measures of motor awareness did not differ significantly ac-
cording to whether subjects were aware of the target shift or
not. This demonstrates the novel finding that motor awareness
can dissociate from perceptual awareness. That is, despite re-
porting that the target did not shift, subjects were neverthe-
less aware that their movement contained a lateral deviation
in the appropriate direction. They are able to report their mo-
tor awareness of their movement despite having no conscious
knowledge of the target event that drove it. This pattern of
results seems quite paradoxical. The subjects clearly knew, in
some sense, that their initial movements contained a lateral
deviation. It is tempting to speculate on how subjects recon-
cile this awareness of their own motor adjustment deviations
 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237 233

with manifest unawareness of any reason why the movement
required adjusting. Subjects did not seem at all troubled by
this incompatibility, and did not seem to find the motor ad-
justments phenomenologically remarkable in the way that
non-intentional movements evoked by transcranial magnetic
stimulation, or by Kohnstamm’s manoeuvre often are.
5. Experiment 2
motor adjustments in our first experiment might not arise in
the parietal structures that generate the adjustment itself, but
in a separate cortical pathway. This reasoning suggested the
intriguing possibility that parietal TMS might suppress the
visuomotor adjustment itself, while leaving the subject with
the (erroneous) conscious awareness that they had adjusted
their movement. Experiment 2 sets out to investigate this pos-
sibility.

5.1. Introduction 6. Experiment 2

Our initial aim in this experiment was to try to dissociate
motor performance from motor awareness. It is widely ac-
knowledged that much, even most, motor function does not
require conscious thought (Haggard, 2001). The term “auto-
matic” is often used in such cases. Therefore, motor perfor-
mance is often dissociated from motor awareness in everyday
life. However, to understand why motor performance some-
times does enter awareness and sometimes does not, an inter-
ventive experimental approach is required, in which the ex-
perimenter can manipulate whether a given feature of move-
ment either does or does not enter awareness. One previous
attempt at this approach was the work of Haggard and Magno
(1999). They used the well-established delay in reaction times
caused by appropriately timed transcranial magnetic stimu-
lation (TMS) over the motor cortex to delay subjects’ motor
performance (Day et al., 1989). Their subjects additionally
judged the perceived time of their reactions, using the method
of Libet and colleagues (Libet, Gleason, Wright, & Pearl,
1983). The results showed that subjects were largely unaware
of the performance delay produced by cortical TMS, suggest-
ing at least a partial dissociation between motor performance
and motor awareness.
More recently, Desmurget et al. (1999) reported that visuo-
motor adjustments in the double-step task could be abolished
by TMS over the posterior parietal cortex (PPC). Moreover,
several authors have additionally proposed that the parietal
cortex acts as an “auto-pilot” for aimed movement (Pisella
et al., 2000) and does not contribute to conscious awareness.
We therefore reasoned that the motor awareness of visuo-
6.1. Method
The method for experiment 2 was identical to that of ex-
periment 1, which in turn was partly based on the study of
Desmurget et al. (1999). Subjects again made reaching move-
ments to targets that were unpredictably perturbed. In this
experiment, however, TMS was delivered on 50% of trials
selected at random. On those trials, TMS was delivered at the
start of movement when the hand velocity exceeded 5 cm/s.
A focal TMS coil was held by the experimenter over the
left PPC. The exact position was localised according to the
method of Desmurget et al. (1999); 4.5 cm caudal and 0.5 cm
medial from the hand motor area. Output was set at 120%
of the subject’s motor threshold. Horizontal EOG was mea-
sured with surface electrodes. Target jumps were triggered at
a random 25–100 ms interval after the onset of EOG activity
associated with the large saccade from peripheral fixation to
the central target. The methods were otherwise as for exper-
iment 1.
Five new right-handed subjects (4 female, mean age 26
years) participated in the experiment.
7. Experiment 2
7.1. Results
Subjects reported perceptual awareness of a target shift on
47% of jump trials in which TMS was delivered, and 47%

Table 2
Mean and S.D. across subjects of endpoint × deviation (mm), for initial and reproduced movements with and without TMS
Position of target Left (target = −40 mm) Centre (target = 0 mm) Right (target = 40 mm)

TMSa No TMSa TMSa No TMSa TMSa No TMSa

Awareb Unawareb Awareb Unawareb n/ab n/ab Awareb Unawareb Awareb Unawareb
Initial movement
Mean (mm) −12 −18 −15 −17 22 21 64 62 62 62
S.D. (mm) 46 39 44 44 32 35 52 50 47 54
Reproduced movement
Mean (mm) −30 −24 −30 −28 18 19 72 67 69 67
S.D. (mm) 36 38 29 36 34 38 45 46 39 52
Note: Negative numbers indication mm left of centre, positive numbers are mm right of centre.
a Presence or absence of TMS.
b Subjective awareness of target jump.
234 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237
Fig. 5. Mean endpoints in perturbation (“jump”) trials (mm).
of jump trials without TMS. A 2.7% of trials were elimi-
nated from the kinematic analysis because of pre-emptive
eye or hand movements, 10.6% because of equipment fail-
ure, and 8.2% because of movements in the wrong direction
with respect to the target shift. The endpoints for initial and
reproduced perturbed movements are shown in Table 2 and
Fig. 5.
We first analysed the no TMS trials to assess whether the
pattern of results found in experiment 1 was replicated in
experiment 2. To establish whether visuomotor adjustments
had been made, the endpoints in each target jump condition
were compared with the endpoints of the unperturbed centre
target trials.
Family wise Bonferroni corrections were performed, and
the significance level for all of the following t-tests is 0.025.
All of the non-TMS jump trials differed significantly from
the unperturbed trials regardless of whether the subject was
aware (left: t4 = 10.85, P < 0.001, right: t4 = −8.69, P = 0.001)
or unaware (left: t4 = 8.30, P = 0.001, right: t4 = −7.19,
P = 0.002) of the target shift. This confirms that visuomo-
tor adjustments were being made regardless of perceptual
awareness.
To establish whether these adjustments entered into mo-
tor awareness, as they had in experiment 1, the reproduced
endpoints of jump trials were compared with the reproduced
endpoints from the centre point condition. In the reproduced
movements, jump trials again differed significantly from un-
perturbed trials regardless of whether the subject was aware
(left: t4 = 6.28, P = 0.003, right: t4 = −10.32, P < 0.001) or un-
aware (left: t4 = 7.84, P = 0.001, right: t4 = −6.56, P = 0.003)
of the target shift.
Finally, to confirm that perceptual awareness had no effect
on motor awareness reproduced endpoints of jump trials in
which subjects were aware of the target shift were compared
with those on trials in which they were unaware. No signif-
icant difference was found for leftward (t4 = 0.39, P = 0.71)
or rightward (t4 = 0.45, P = 0.68) adjustments. The pattern of
results seen in experiment 1 was therefore replicated in the
non-TMS trials of experiment 2.
7.1.1. No effect of TMS on visuomotor adjustment
A further series of comparisons focussed on effects of
TMS on visuomotor adjustment. The endpoints of the TMS
jump trials were compared with the endpoints of the unper-
turbed TMS centre point trials. Family wise Bonferroni cor-
rections were again performed, and the significance level for
all of the following t-tests is 0.025.
All of the TMS jump trials differed significantly from the
unperturbed TMS trials regardless of whether the subject was
aware (left: t4 = 7.88, P = 0.001, right: t4 = −6.08, P = 0.004)
or unaware (left: t4 = 8.90, P = 0.001, right: t4 = −7.51,
P = 0.002) of the target shift. Therefore, visuomotor adjust-
ments were clearly present in trials where TMS was applied
to the PPC.
Although TMS did not abolish visuomotor adjustments
it might nevertheless have subtly changed their kinematic
pattern. Therefore, the initial movement endpoints of jump
trials in which TMS was administered were compared with
the endpoints of no TMS jump trials. No significant differ-
ences were found regardless of whether subjects were aware
(left: t4 = −1.87, P = 0.14, right: t4 = −0.38, P = 0.73) or un-
aware (left: t4 = 0.24, P = 0.82, right: t4 = 0.19, P = 0.86) of
the target shift.
We also speculated that TMS could have been effective in
just a small subset of trials. However, plotting the endpoints
for individual trials showed essentially overlapping endpoint
distributions for TMS and non-TMS trials (Fig. 6).
Finally, although it would have been contrary to our pre-
dictions, it could still have been possible that TMS had af-
fected motor awareness, despite motor performance remain-
ing unaffected. Therefore, the reproduced endpoints of the
TMS jump trials were compared with their non-TMS coun-
terparts. No significant differences were found (all t4 < 2, all
P > 0.20).
8. Experiment 2
8.1. Discussion
Three features of experiment 2 are noteworthy. First, we
replicated in a further group of subjects the dissociation
between motor awareness and perceptual awareness seen in
experiment 1. Again, we observed that subjects reproduced
motor adjustments following target jumps of which they were
perceptually unaware. The use of EOG to trigger the target
jump in the present experiment is a methodological improve-
ment which makes the lack of any perceptual awareness
effects more convincing, if anything, than in experiment 1.
Second, we entirely failed to obtain the suppression
of visuomotor adjustments by parietal TMS reported by
 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237 235

Fig. 6. Endpoints of perturbed and unperturbed trials for each subject, with and without TMS (mm). The target was located at (0, 240).

Desmurget et al. (1999) despite using comparable methods.
We speculate on two possible reasons for this. Although we
used their procedure for localising the TMS site, Desmurget
et al. subsequently confirmed the location of their stimula-
tion using MRI. They also noted that one of their subjects
who showed a slightly different pattern of results had been
stimulated more medially than the others. We did not con-
firm our stimulation site with MRI, so anatomical variations
or incorrect localisation might have resulted in inappropriate
stimulation sites for some of our subjects. On the other hand,
one of their subjects who did not show the inhibitory effect
of TMS, did appear to have been stimulated appropriately.
Therefore, there may be substantial individual differences in
responsiveness to parietal TMS. Finally, the TMS coil and
stimulator in their study may have been more powerful than
ours. In particular their stimulation may have been effective
at deeper sites within the intraparietal sulcus than ours.
Third, we found no effects of parietal TMS on motor
awareness, measured as the discrepancy between initial and
reproduced visuomotor adjustments. This is, of course, a null
result, and we had not predicted any such effects. Moreover,
one might argue that the absence of any TMS effect on motor
performance would make effects on motor awareness un-
likely. Nevertheless, we note that our failure to alter motor
awareness is at least consistent with the view that the PPC
provides an “auto-pilot” for on-line movement control, (Gréa
et al., 2002; Pisella et al., 2000) independent of conscious ex-
perience.
8.2. General discussion and conclusions
Previous studies have suggested that the motor system can
respond to information of which we are perceptually unaware
(Pélisson et al., 1986) Our own work had further shown that
236 H. Johnson, P. Haggard / Neuropsychologia 43 (2005) 227–237
people’s awareness of their own motor output may involve
an attenuated or an exaggerated perception of their actual
movement, essentially depending on the extent to which the
movement was intended or not (Johnson et al., 2002). In the
present study, we have investigated the link between these two
approaches, by investigating whether perceptual awareness of
a target shift modulates motor awareness of the visuomotor
adjustment to that shift during reaching movements.
Our results show that awareness of adjustment to ongoing
movements is independent of perceptual awareness of any tar-
get displacement necessitating such adjustment. Therefore,
we have shown that motor awareness and perceptual aware-
ness can be dissociated, at least in the double-step task. This
dissociation appears relatively complete: the magnitudes of
lateral deviations in reproduced movements were numeri-
cally close to those in the corresponding initial movements,
and not just attenuated versions of the original movements. To
conclude, we comment on some psychologically important
features of this dissociation.
Our result suggests a paradox or incompatibility at the
heart of subject’s experience of their own action. In some tri-
als, our subjects reported a stationary target on the one hand,
yet a clearly deviated trajectory on the other. At face value,
resolution of these incompatible situations would seem possi-
ble only if subjects generate an alternative explanation, other
than a displacement of the target, of why they made a tra-
jectory deviation. For example, subjects could conceivably
attribute the deviation of their initial movement to an execu-
tion error that accidentally arose in the course of a movement
to an apparently stationary target. They would then faith-
fully reproduce this error. Two reasons lead us to reject this
idea. First, our subjects did not show the sort of frustration
and expletives typically associated with the experience of
movement error. Second, subjects presumably knew that their
lateral deviations were larger than typical execution errors
(van Beers, Haggard, & Wolpert, 2004). Instead, we think
the mind does not even attempt to resolve lack of percep-
tual awareness with positive motor awareness. Rather, motor
awareness and perceptual awareness might be quite different
and independent processes in the brain, which have different
neural substrates and different natural time courses. In some
respects, this position recalls the concept of multiple “micro-
consciousnesses” within the visual system (Zeki & Bartels,
1999).
We end by noting a possible relation between our con-
cept of motor awareness and the concept of agency used by
Jeannerod (2002). The sense of agency derives from an ability
to compare motor commands with reafference from the body
movements and external events caused by the commands.
Our data suggests that subjects can have a clear conscious
awareness of their own movement patterns even when the
external stimulus event causing the movement is unclear. We
believe the dissociation between perceptual awareness and
motor awareness may be an important clue towards under-
standing how the brain distinguishes between self-generated
and external events, and towards understanding both the
brain processes and the conscious experience of being an
agent.
Acknowledgements
HJ was supported by a BBSRC studentship. Further sup-
port was provided from a BBSRC Project grant to PH, a Lev-
erhulme Research Fellowship awarded to PH, and a Royal
Society Equipment grant.We are grateful to Louise Whiteley
for assistance with preparing this work for publication.
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