~

Pergamon

0028 3932(95)00128-X

Neuropsycholoyia,Vol. 34, No. 5, pp. 369-376, 1996

Copyright ~ 1996 Elsevier Science Ltd. All rights reserved
Printed in Great Britain
0028 3932/96 $15.00+0.00

Visual illusion and action

MAURIZIO

GENTILUCCI*,
M. CRISTINA

SERGIO CHIEFFI,
SAETTI and IVAN

ELENA
TONI

DAPRATI,

lstituto di Fisiologia Umana, Universit~ di Parma, via Gramsci 14, 1-43100 Parma, Italy
(Received 11 March

1995;

accepted 2 Auyust

1995)

Abstract--The role of allocentric cues on movement control was investigated in the present study. Pointing movements directed to
the more distant vertex of closed and open configurations of the M~iller-Lyer illusion, as well as to the vertex of control lines, were
studied in four experimental conditions. In the first (full-vision condition) subjects saw both stimulus and their hand before and
during movement, in the second (non-visual feedback condition) they saw the stimulus, but not their hand during movement. In the
two remaining conditions (no-vision conditions) vision of the scene and the hand was precluded. Pointing was executed 0 sec (no
vision 0 sec delay condition) or 5 sec (no-vision 5 sec delay condition) after the light was switched off. The Mi~ller-Lyer illusion
affected pointing kinematics with respect to the control lines. Subjects undershot and overshot the vertex location, respectively, of
the closed and open configuration. Correspondingly, the entire kinematics were changed. The main result was, however, a gradually
increasing effect of the perceptual illusion when pointing was executed from memory compared to the full-vision condition. These
data are discussed according to the hypothesis that the system underlying visual perception in the allocentric flame of reference and
that involved in motor action can functionally interact. The strength of this interaction depends upon the efficiency of the egocentric
flame of reference by which motor actions are constructed.

Key Words: Miiller-Lyer illusion; pointing; humans; visual feedback; non visual feedback; visual memory.

had the strong illusion that the target had changed

position. Despite this illusion, subjects directed their saccades to the true location of the target. Gentilucci and
Negrotti [13] observed a different type of constant error
when subjects were required to reproduce a distance
either in a perceptual or in a m o t o r task.
The distinction between the use of either an egocentric
or an allocentric frame of reference for visual analysis of
a scene has its counterpart in dissociation in the visual
cortex between processes involved in perception and
visuomotor integration, as supported by the neuropsychological studies of Goodale et al. [19] and Milner
et al. [25]. The agnosic patient they described showed a
clear deficit in reproducing with her hand either the size
or the orientation of visually presented objects, whereas
her grip size while reaching was normally scaled as a
function of object size [21] and the orientation of her
wrist was appropriate to that of the target in a 'posting'
task.
Efficiency of the egocentric frame of reference is
reduced in some visuomotor tasks, such as in movements
executed without visual feedback or in memory-driven
movements. In the former movements, kinesthetic information, but not visual information about hand position
is available to control movement execution (for the role

Introduction

A pointing movement directed to a target requires calculation of the target's position in space with respect to
the body (an egocentric frame of reference)
[11,27, 31,35]. Encoding the position of a visual target
can be influenced by its spatial relation to surrounding
visual cues (an allocentric frame of reference) [2, 6, 12, 34].
Information in an allocentric frame of reference is necessarily processed for scene perception [23]. It has been
shown that when the two frames provide conflicting
information such as a misleading perceptual effect, information from the egocentric system is selected for m o t o r
planning. Bridgeman and coworkers [3, 4] have shown
that even if a fixed visual target surrounded by a moving
frame appeared to drift in a direction opposite to that of
the frame, subjects continued to point to the true location
of the target. Wong and Mack [37] have found that, when
a target was re-presented after a 500 msec blank interval
in the same location, but with the surrounding frame
displaced a few degrees to the right or to the left, subjects

* To whom correspondence should be addressed; fax: 39-521291304.

369

370

M. Gentilucci et al./Illusion and action

o f p r o p r i o c e p t i v e i n f o r m a t i o n on m o t o r c o n t r o l see G e n tilucci et al. [14]). In the latter m o v e m e n t s , c o n t r o l is

based, in a d d i t i o n , on stored i n f o r m a t i o n a b o u t object
properties. In b o t h cases, i n f o r m a t i o n f r o m the allocentric ( ' p e r c e p t u a l ' ) frame m a y be e m p l o y e d to guide
the m o v e m e n t to the target. To s u p p o r t this hypothesis,
G n a d t et al. [17] observed t h a t b o t h errors a n d kinematics
o f saccades were different when they were directed either
to a visual or to m e m o r i s e d target [17]. A b s o l u t e errors
increased when p o i n t i n g to a m e m o r i s e d target was
executed with a delay o f m o r e t h a n 2 sec [9].
In the present e x p e r i m e n t we tested the hypothesis t h a t
a g r a d u a l shift exists t o w a r d s the use o f an allocentric
frame o f reference when the possibility to efficiently use
the egocentric frame o f reference c o r r e s p o n d i n g l y
decreases. W e studied p o i n t i n g m o v e m e n t s directed to
targets whose p o s i t i o n in space was e r r o n e o u s l y perceived
because o f an illusion. The M ~ l l e r - L y e r illusion [26] was
used. M o v e m e n t s were executed in c o n d i t i o n s o f full
vision, non-visual feedback, a n d in two no-vision conditions (i.e. f r o m visual m e m o r y ) , n a m e l y 0 sec delay a n d
5 sec delay conditions. It can be assumed t h a t in the first
no-vision c o n d i t i o n , only execution a n d in the second
one b o t h p l a n n i n g a n d execution rely on visual m e m o r y .
A n increasing effect o f the perceptual illusion was
o b s e r v e d when p o i n t i n g was executed f r o m m e m o r y c o m p a r e d to the full-vision condition. These d a t a suggest that
the system underlying visual p e r c e p t i o n in the allocentric
frame o f reference a n d that involved in m o t o r action m a y
functionally interact a n d the strength o f this i n t e r a c t i o n
d e p e n d s u p o n the efficiency o f the egocentric frame o f
reference.

Methods
Subjects
Thirty-two right-handed subjects participated in the present
study. All were naive as to the purpose of the experiment.

Apparatus and procedure

The subjects were seated comfortably, in front of a table and
placed their head on a head-and-chin rest. Their right hand
rested on the plane of the table and their right index finger was
positioned on a disk (starting disk, Fig. 1). The starting disk (1
cm diameter) was along the subject's sagittal plane.
Visual stimuli were the open and closed configuration of the
Mt~ller-Lyer illusion as well as control lines. Figure 1 shows
both the illusory and control stimuli. Lines were of three different lengths (8, 12, and 16 cm). The ratio between lengths of
wings and shafts was 0.47. The inclination of wings with respect
to direction of shaft was 45 . Stimuli were drawn in black ink
on white panels which could be either placed on the table in
front of the subject or projected on the plane of the table by
means of a semi-reflecting mirror. In this latter condition (nonvisual feedback condition, see below) an apparatus was used
that allowed vision of the stimulus, but not of the hand (for a
description of the apparatus, see Gentilucci et al. [14]). The

main axis of the stimulus was approximately aligned with the

subjects' body midline. The starting disk was located at 1 cm
from the proximal vertex of each stimulus line. Thus, the position of the starting disk varied from trial to trial as a function
of the length of the visual stimulus (20 cm from the subject's
frontal plane for the 8 cm line, 18 cm for the 12 cm line, and 16
cm for the 16 cm line).
Subjects were required to execute a pointing movement from
the starting disk to the more distant vertex of the stimulus. The
verbal instruction was: "Point to the conjunction of the three
farther lines, neglecting the surrounding scene". Subjects were
required to perform the movement as naturally as possible, with
the same velocity as during spontaneous movements.
Pointing movements were executed in four experimental conditions: full-vision condition, non-visual feedback condition,
no-vision 0 sec delay condition and no-vision 5 sec delay
condition. In all conditions, at the beginning of each trial,
subjects were free to inspect the panel for 5 sec. In the fullvision condition, light in the room remained on over the whole
course of the trial. The go signal was given by the experimenter,
5 sec after presentation of the stimulus. Subjects could see both
their hand and the target, before and during movement. In the
non-visual feedback condition subjects saw the stimulus, but
not their hand during movement. They could see their hand
only before movement. Instruction to move was given as in the
previous condition. In the no-vision 0 sec delay condition, after
the 5 sec inspection period, switching off the light was the signal
for the subjects to start the movement. In this condition subjects
saw neither stimulus nor their hand during movement. Finally,
in the no-vision 5 sec delay condition, the go signal was given
by the experimenter 5 sec after the light was switched off.
The 32 subjects were divided into four groups. Each group
was tested in one of the four conditions. The group tested in
the no-vision 5 sec delay condition was constituted of five
females and three males. The three other groups were composed
of five males and three females. The age of the subjects ranged
from 18-31 yr. An analysis of variance (ANOVA) in which the
dependent variable was the age of the subjects, showed no
statistical difference between the groups [ F ( 3 , 2 8 ) = 0.68,
P = 0.57].
In each experimental condition, 72 trials were run; that is
eight repetitions of each combination of line length and stimulus
configuration. Order of presentation of both stimulus configurations and line lengths was randomised.

Movement recording and data analysis

Pointing movements were recorded for subsequent analysis
with the opto-electronic ELITE system (B.T.S., Milan, Italy).
This system, consisting of two cameras and a processor, detects
spatial position of infrared reflecting markers. Apparatus and
data analysis procedure are described elsewhere [14]. In the
present study two markers were used. The first marker was
placed on the tip of the subject's right index finger. The second
marker was placed on the table along the subject's sagittal plane
at 7 cm from the edge of the table. It was used as a reference
point (centre of the coordinate system).
The marker placed on the tip of the subject's right index
finger was used to analyse pointing kinematics. The procedure
to calculate the beginning and the end of the movement is
described elsewhere [14]. Trajectories were normalised in time,
that is equal numbers of points describing trajectories in space
were computed (40 frames, each lasting 20 msec). After that,
mean trajectories of each subject for each condition on the
horizontal plane were calculated. In the present study the following parameters were analysed: movement amplitude, component of movement amplitude along the subject's transverse
axis (final deviation in the transverse axis), movement time,

M. Gentilucci et al./Illusion and action

0$

371

o$

Fig. 1. The open (right position) and closed (left position) configurations of the M011er-Lyer illusion and the control line (middle
position). S: starting position.

peak velocity and time to peak velocity. The standard deviations

of final finger position on the subject's sagittal and horizontal
axis were also calculated. The square root of the sum of their
squares was used as a measure of variability of final finger
position (final variability on the horizontal plane).
The kinematic parameters were submitted to ANOVAs
whose between-subjects factor was experimental condition (fullvision, non-visual feedback, 0 sec delay, 5 sec delay) and withinsubjects factors were length (8, 12 and 16 cm) and configuration
(open, closed and control). The Newman-Keuls test was used
as a post-hoe test. Further ANOVAs were carried out in which
the dependent variables were the differences between movement
amplitudes measured for each of the two illusory configurations
and movement amplitude measured for the control configuration. The between-subjects factor was condition (full-vision,
non-visual feedback, no-vision 0 sec delay, no-vision 5 sec
delay), and the within-subjects factors were length (8, 12 and 16
cm) and configuration (open and closed configuration).

in non-visual feedback conditions where only the open

configuration was effective with respect to the closed and

Stimuluslength:8 cm

,2o
1

Full
vision

Non
visual
feedback

0-sec
delay

5-see
delay

Stimuluslength:12cm

1 Closed

Results
Figure 2 shows movement amplitudes averaged across
the subjects in the various experimental conditions. N o t e
an effect of the illusory context in all conditions. In the
A N O V A m o v e m e n t amplitude varied significantly with
both illusory configurations with respect to the control
lines [F(2, 56) = 82.9,P < 0.00001]. Subjects undershot
the vertex o f the closed configuration (124.2 mm) and
overshot that o f the open configuration (130.2 ram), with
respect to the control lines (125.8 mm) [P < 0.05]. Experimental condition was significant [F(3.28) = 4.4,
P < 0.01; full vision: 129.0 mm, non-visual feedback:
138.8 mm, no-vision 0 sec delay: 121.0 ram, no-vision 5
sec delay: 118.3 mm]. Post-hoc comparison showed one
significant difference between the non-visual feedback
and the two m e m o r y conditions [P < 0.05].
The main result was that the illusion effect differed
a m o n g the four experimental conditions (interaction
between configuration and experimental condition,
[F(6, 56) = 8.0,P < 0.00001], see Table 1 for numerical
values). The smallest effect was f o u n d in full-vision and

[] Control
[] Open

Full
vision

Non
visual
feedback

O-sec
delay

5-sec
delay

Stimuluslength:16cm

2oo

/
Full

Non

vision

visual
feedback

0-sec
delay

5-sec
delay

Fig. 2. Mean values of the movement amplitude in the four

experimental conditions measured for the three stimulus lengths
and configurations. Note that the starting position was displaced 1 cm from the proximal vertex of the stimulus. Since
movement amplitude was calculated from this position, its value
was longer than stimulus length, ms, Closed; U], control;
[], open.

M. Gentilucci et al./lllusion and action

372

Table 1. Mean values (SE) of movement spatial parameters

Full vision

Non-visual
feedback

0 sec delay

5 sec delay

Movement
amplitude
(ram)

Closed
Control
Open

128.0 (0.7)
128.4 (0.6)
130.4 (0.6)

136.8(4.2)
137.9(4.2)
141.6 (4.3)

118.5(4.5)
120.4(4.4)
124.0(4.8)

113.6(6.4)
116.5(6.6)
124.8 (7.0)

Final deviation
in the transverse axis
(turn)

Closed
Control
Open

3.9 (0.2)
3.7 (0.3)
3.8 (0.4)

-0.6(1.6)
-0.9(1.8)
-2.6(1.7)

0.8(1.7)
0.0(1.9)
0.2(1.9)

-0.3(0.8)
1.5(1.0)
1.8(1.1)

Final variability on
the horizontal plane
(mm)

Closed
Control
Open

3.9 (0.4)
3.3 (0.1)
3.6 (0.2)

4.9(0.9)
4.8(1.0)
4.3 (1.2)

9.2(0.5)
9.7(0.6)
10.1 (0.6)

14.8(0.8)
13.8(0.8)
15.9(1.0)

the control configurations [P < 0.05]. The illusion effect

increased in the two memory conditions in which both
the open and the closed configurations had an influence
on movement amplitude [P < 0.05].
The ANOVA in which the dependent variable was
the variation of the movement amplitude in the open
configuration condition with respect to the control condition (see Methods) showed that the largest modification
was found in the no-vision 5 sec delay condition. The
increase in movement amplitude was significantly greater
in this condition than in the other three experimental
conditions [F(3, 28) = 13.4,P < 0.00005, 8.3 mm vs 3.6,
3.7, 2.1 mm, P < 0.0005]. In summary, the illusion effect
increased progressively from the visual conditions to the
memory conditions, being more evident for the open
configuration.
Stimulus length was also a variable in inducing the
illusion effect, which appeared to increase with stimulus
length (interaction between stimulus length and configuration, F(4, 112) = 3.1, P < 0.05]. This was particularly so for the closed configuration. Indeed, it
induced a significant variation in movement amplitude
only when stimulus length was 16 cm [P < 0.0005].
Figure 3 shows the mean trajectories of four subjects
on the horizontal plane executed in the various
conditions. The reported variations in movement amplitude following presentation of the Mtiller-Lyer configurations could be due to selective lateral trajectory
deviations and/or changes in final trajectory variability.
This could depend on the different wing orientations of
the figures, inducing more or less straight hand paths.
This possibility was tested by taking into account final

deviation in the transverse axis and final variability on the

horizontalplane (see Table 1). These parameters were not
influenced by the illusion. Final deviation in the transverse axis showed one significant interaction between
experimental condition and configuration [F(6,56)-2.89, P < 0.05]. A significant drift to the left was observed
for the open configuration in the non-visual feedback
condition [P < 0.05]. As expected [14] final variability on
the horizontal plane was significantly affected by experimental condition [ F ( 3 , 2 8 ) : 28.2, P < 0.00001, vision,
3.6 mm, non-visual feedback, 4.7 mm, no-vision 0 sec

delay, 9.77 ram, no-vision 5 sec delay, 14.8 mm]. In

addition, there was a significant increase of variability
for each increase of stimulus length [ F ( 2 , 5 6 ) = 24.0,
P < 0.00001, 8.4 mm vs 10.0 mm vs 11.1 mm, post-hoc
test, P < 0.05]. This increase was more evident in the two
memory conditions (interaction between experimental
condition and
stimulus length:
[F(6, 56) = 5.1,
P < 0.0005].
Table 2 shows the main kinematic parameters describing pointing trajectory. Movement time increased when
pointing was executed to the vertex of the open configuration (668.2 msec) with respect to both control line
(648.2 msec) and closed configuration (642.4 msec)
[F(2, 56) = 17.3, P < O.O0001,post-hoe test: P < 0.0005].
Variations in movement time with both illusions showed
a trend to differ among the four experimental conditions
(interaction between experimental condition and configuration: IF(6, 5 6 ) = 2.3, P = 0.052]). The open configuration induced a significant variation only in the two
memory conditions [P < 0.05], whereas the closed configuration modified movement time only in 5 sec delay
condition [P < 0.05]. Movement time increased significantly for each increase in stimulus length [F(2, 56) = 162.7,
P < 0.00001, 590.0 msec vs 660.0 msec vs 708.0 msec,
post-hoc test, P < 0.0005].
Peak velocity [ F ( 2 , 5 6 ) = 5.1, P < 0.01] and time to
peak velocity [F(2, 56) = 3.8, P < 0.05] were influenced
only by the open configuration (peak velocity, 420.6
mm/sec vs 411.8 and 412.7 mm/sec, post-hoc test,
P < 0.05; time to peak velocity, 271.0 msec vs 239.8 and
258.9 msec, post-hoc test, P < 0.05). Length significantly
affected peak velocity [ F ( 2 , 5 6 ) = 581.6, P < 0.00001]
and time to peak velocity [F(2, 56) = 14.2, P < 0.00001].
That is, peak velocity (323.9 mm/sec vs 415.7 mm/sec vs
505.5 mm/sec,post-hoe test, P < 0.0005) and time to peak
velocity (240.0 msec vs 258.0 msec vs 270 msec, post-hoc,
test, P < 0.05) increased significantly for each increase in
length.
Discussion

The present study investigated the possibility that target position coded in an allocentric frame of reference

M. Gentilucci et al./Illusion and action

200

200

373

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-50

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loo

200

50

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-50

50

200
2

3
100

-50
200

32
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100

i
0

100

50

-50

200

50

-50

200

50

i
3

lOO

o
-50
200

31

I00

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50

I00

0
-50
200

0
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200

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12
1

3
100

i00

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50

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100

mm

50

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50

Fig. 3. Representative mean trajectories on the horizontal plane of four subjects. Trajectories were averaged across seven trials after
normalisation in time (40 frames). Each row shows trajectories of one subject. Moving from top row to bottom row movements
executed in visual, non-visual feedback, no-vision 0 sec delay, no-vision 5 sec delay conditions are presented, respectively. Left,
middle, right row shows movements executed when stimulus length was 8, 12 and 16 cm, respectively. Trajectory 1,2, 3 refers to
when closed configuration, control line, and open configuration were, respectively, presented. Abscissa axis: transverse axis, ordinate
axis: sagittal axis. The origin of the axes corresponds to the hand starting position. Plus sign was used for movements to the right,
minus sign for movements to the left.

gradually influences v i s u o m o t o r transformation when

the efficiency o f the egocentric frame o f reference correspondingly decreases. To this purpose, subjects pointed
to a target whose position could be erroneously localised
because o f an illusory effect. The Mtiller-Lyer illusion was
used. Movements were executed in different experimental
conditions in which the possibility o f an efficient comparison between target and h a n d position gradually
decreased. D u r i n g movements executed without visual
feedback h a n d vision was precluded. In no-vision conditions only stored information o f target position was

used to guide movements (0 sec delay) and also to plan

them (5 sec delay). Consequently, in each of the four
conditions information about target position could be
differently processed in order to execute an appropriate
movement.
The illusion affected movements in all conditions.
Overshooting was observed with the open configuration,
undershooting was observed with the closed configuration. This became more evident with increasing stimulus
length. Illusion effects f o u n d in previous experiments
[10, 15,29, 30] were greater in percentage than in our

374

M. Gentilucci et al./lllusion and action

Table 2. Mean values (SE) of movement kinematic parameters
Full vision

Non-visual
feedback

0 sec delay

5 sec delay

Movement
time
(ms)

Closed
Control
Open

641.2(30.9) 661.1(37.1)
645.4(27.5) 646.3(33.2)
657.6(31.9) 671.3(35.4)

653.3(33.7)
668.6(40.3)
682.4(36.4)

614.4(33.5)
632.5(41.0)
661.7(39.1)

Peak
velocity
(ram/s)

Closed
Control
Open

441.1 (13.1) 427.3(27.2) 384.1(20.9)

442.0(8.9) 428.2(27.9) 381.6(23.3)
439.9(15.6) 434.0(27.9) 393.7(27.1)

394.8(27.3)
398.8(33.1)
414.8(26.4)

Time to
peak velocity

Closed
Control
Open

208.3 (16.2) 285.7(27.8)

208.9(16.1) 281.4(28.4)
221.6(20.0) 291.6(29.4)

252.1(24.9)
242.1(19.7)
249.6(20.9)

experiment. This difference is likely due to the different

instruction given to subjects. In the previous experiments
they were required to judge a length, whereas in our study
explicit instruction was given to localise the configuration
vertex, neglecting the surrounding figure. The open configuration caused a greater effect than the closed configuration. This is consistent with the result that
perceptual judgement about shaft length is more influenced by the open than by the closed configuration [22].
The Mtiller-Lyer illusion affected not only movement
amplitude, but also the entire movement kinematics, that
is peak velocity, time to peak velocity, and movement
time. Consequently, the illusion affected the early phase
and perhaps even the planning of the movement. This
finding seems to be at odds with the results of Mack
et al's. experiment [22] in which no illusion effect was
observed for the same type of response. In that experiment, however, stimuli were transversally located. In our
experiment they were sagittally located and consequently
they could influence hand trajectory, Indeed, results of
previous experiments [28, 32] indicate that visual stimuli
placed in the space between hand starting position and
target are more relevant than stimuli located elsewhere.
In addition, in Mack et al's. experiment [22] subjects had
to fixate either the vertex of the configuration or another
point in space, whereas in the present experiment they
were free to inspect the configuration. It has been proposed that efferent copy of eye movements during inspection of MOller-Lyer configurations can contribute to
increase the illusion effect [10].
The main result of the present experiment was that
illusion differently affected movement amplitude across
the different experimental conditions. That is, the effect
was very small (2 ram) in the full-vision condition and
gradually increased moving from the non-visual feedback
condition to the two memory conditions. The possibility
that the different wing orientations presented in the three
configurations induced different lateral deviations of
movement end point was examined. The open configuration could have induced more dispersion of trajectories than the closed configuration because its wings
did not interfere with the hand path. This effect became
evident when visual feedback corrections were not poss-

275.6(44.9)
269.3(39.5)
286.6(44.8)

ible. Thus, lateral deviation, rather than the illusion

effect, might be responsible for variations in movement
amplitude. However, this was not the case. First, final
deviation in the transverse axis and final variability were
not influenced by the illusion. Final variability increased
only in the non-visual conditions and with distance, as
found in a previous experiment [14].
The finding that memory-driven movements were
influenced by perceptual coding of the target is in agreement with the results by Goodale et al. [18]. However, in
their experiment a more abstract object representation
was used because movement had to be pantomimed and
consequently imagined. On the contrary, in our experiment movement had to be directed at the real target.
Another difference between their results and ours is the
following: our data suggest the existence of a gradual
crosstalking between 'perceptual' and 'pragmatic' object
representations. This occurred when the efficiency of the
egocentric frame of reference was correspondingly
reduced.
The problem arises as to how end point is coded when
comparison between target and body reference gradually
becomes difficult. A first possibility is that prime information on the end point is coded in an allocentric frame
of reference. That is, the relationships between target
and surrounding objects are primarily analysed. In our
experiment, localisation of the vertices of the MtillerLyer figures was therefore affected by structural factors
[1,5, 16, 36] and cognitive mechanisms [7, 8, 20] inducing
illusion effects. Movement execution needs, however,
translation of the end point into an egocentric frame of
reference. Selection mechanisms active in the egocentric
system allow to focus on the area around the distal vertex
of the configuration. The consequent exclusion of the
proximal part of the figure reduces the illusion, likely
inactivating its cognitive component. This mechanism
can account for previous findings [3, 4, 37] showing a
dissociation between perceptual judgement and motor
output. The increasing effects of the illusion in the memory conditions can depend on a progressive decay in the
working memory of the selection mechanism's efficiency.
An alternative possibility is that movement end point
is simultaneously coded in both egocentric and allocentric

M. Gentilucci et al./lllusion and action

frames of reference. In the egocentric system the correct
end point is extracted from the background and referred
to the body. In the allocentric frame of reference the same
point in space is considered, but as a part of the illusory
context. That is, it is displaced according to the presented
configuration. Information from both reference frames is
used in order to plan movement. Interaction between the
two frames increases when the availability of efficiently
using egocentric cues is reduced, as it happens in movements either without visual feedback or driven by visual
memory. This hypothesis could be in contradiction with
the finding that an effect of illusion, although small, was
observed in the full-vision condition, when the egocentric
frame should be highly efficient. However this effect is
likely explained by the optical and retinal component of
the illusion [5], that is by that component occurring in
the early visual pathway.
In conclusion, the first hypothesis suggests a cascade
construction in which space is built according to sequential object representations, whereas the second suggests
two parallel constructions of space in which multiple
representations are possible. The two hypotheses can be
accounted for by two theories about the processes performed in the dorsal and the ventral visual pathways.
The first theory [33] proposes that space is unitarily constructed in the dorsal visual stream ('where pathway'),
whereas object features are computed in the ventral visual
stream ('what pathway'). The second theory [19, 24] suggests that the same object properties are represented in
parallel in the two visual pathways. The dorsal pathway
is involved in "how' to interact with the object (a visuomotor representation), the ventral one in elaborating
'what' the object is (a perceptual representation). Accordingly, two parallel representations of space are suggested:
one coded in egocentric frame of reference in the dorsal
visual stream, the other in allocentric frame of reference
in the ventral one. Our data indicate that the allocentric
representation concurs with, or, in other words, tunes
space representation for motor output.

3.

4.

5.

6.

7.

8.

9.

10.

11.

12.

13.

14.

15.

Acknowledgements~--The authors are grateful to Dr D. Carey

for valuable comments on the manuscript; they thank Prof A.
Allport, Dr M. Gangitano and Dr L. Fadiga for discussion of
data. The work was supported by a Research grant from the
Human Frontier Science Program, and by grants CNR and
MURST to M. G.

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