Neuropsychologia,Vol. 35, No. 12, pp.

1577-1582, 1997

Pergamon

~'; 1997 Elsevier Science Ltd. All rights reserved

Printed in Great Britain
0028 3932/97 $17.00+0.00

PII: S0028 3932(97)00061M

Grasping an illusion
ELENA

DAPRATI

and MAURIZIO

GENTILUCCI]"

Istituto di Fisiologia Umana, Universitfi di Parma, via Gramsci 14, 1-43100 Parma, Italy

(Received 23 July 1996; accepted 22 April 1997)

Abstract--In the present study we attempted to determine the nature of the visual analysis that is performed on an object in order

to grasp it. We required eight healthy subjects to reach and grasp a wooden bar which was superimposed over the shaft of the
Mt~ller-Lyer illusion. Vision of both the hand and the bar was allowed. Three different bar lengths were used. Two additional control
tasks in which the subjects were required to reproduce the length of the shafts were carried out. The results showed that hand shaping
while grasping the bar was influenced by the illusion configurations on which it was superimposed. However, this effect was smaller
than that observed in the two tasks of length reproduction. These results support the notion that visual analysis performed on the
object of a grasp movement is global and takes into account the object itself, as well as its relationships with surrounding cues. We
propose, as suggested previously for reaching movements (Gentilucci, M. et al., Neuropsychologia, 1996, 34, 369-376), two partially
independent stages during visuo-motor integration for grasping an object. In the first stage, the object is coded inside an objectcentred frame of reference. In the second stage it is transposed in an egocentric frame of reference, in which the spatial relations
between object and agent are computed. In this second stage the influence of cues surrounding the target is minimized. 1997
Elsevier Science Ltd. All rights reserved
Key Words: reaching to grasp; Mtiller-Lyer illusion; grasping task; drawing task; matching task; kinematics; healthy humans.

grasping [4, 7, 13, 15, 17, 25]. Until recently, however,

the question as to what processes are used to visually
analyse an object in order to grasp it remains an open
issue. It seems plausible that the first aim of the visual
analysis is the selection of the appropriate opposition axis
for the fingers on the object [12]. We can assume that, at
first, by means of a rough analysis of the space occupied
by the object, the opposition axis aligned along the initial
finger orientation is selected. A more detailed analysis of
the corresponding portions of the object's surface is then
performed in order to determine whether a stable grasp is
allowed. When this does not occur (e.g., the two opposite
surfaces are not parallel), the adjacent opposition axes
and the corresponding object's surfaces are progressively
explored, until the appropriate points are located. Their
position is computed in three-dimensional space with
respect to the body (egocentric frame of reference) and
grasping is achieved by carrying each finger to the appropriate contact point on the target object. According to
this hypothesis, a global visual analysis of the object does
not necessarily take place.
Alternatively, we can suggest that a global visual analysis is performed on the object in order to grasp it. We
define as global a process that implies visual analysis of
the whole object, and the visual relations between the
object and the surrounding scene. It is possible that information extracted from the knowledge of object's use and

Introduction

Although we perform it daily, reaching to grasp an object

(prehension) is by no means a simple m o t o r act. For this
m o t o r act to be successful, the hand must be carried to
the appropriate location in space, and the fingers must
be shaped according to the orientation, size and shape of
the target object. On the basis of behavioural studies
[14, 15] the act of prehension can be divided in two
components: the transport (or reach) and the grasp. It
has been hypothesized that, whereas the transport component is more concerned with visual computation of the
spatial relationships between the target object and the
body, the grasp component depends on intrinsic object
properties, such as size and shape [2, 15]. Independent
support for this hypothesis has come from single-neuron
recording studies in the premotor [6, 22] and parietal
cortices [23, 24].
In particular, the dependence of grasp planning on
visual analysis of intrinsic object properties is demonstrated by numerous observations of a strong correlation between hand shaping and object size during

t Address for correspondence: Istituto di Fisiologia Umana,

via Gramsci 14, 1-43100 Parma, Italy; fax: 521 291 304; e-mail:
fisioum@synet.symbolic.pr.it.
1577

1578

E. Daprati and M. Gentilucci/Grasping an illusion

from relevant semantic features are also involved in this

process. Accordingly, the object is primarily analysed
inside an object-centred frame of reference, independently of the spatial relations between object and
observer.
In previous studies we attempted to determine the nature of the visual analysis used in order to localize the
target of pointing movements. We reported that when
asked to point to the distal vertex of the Mfiller-Lyer
illusion [19], subjects misplaced the target according to
the presented configuration. This effect was present in
movements executed under visual control and it increased
for memory-driven movements, as well as in conditions
by which the efficiency of the egocentric system was
reduced [8]. We argued that subjects acquired information on target location by means of a global analysis
of the figure in allocentric coordinates (as demonstrated
by the illusion effectiveness), and successively transposed
the target in egocentric coordinates (as suggested by the
modulation of the illusion effect).
It has been shown previously that the visual analysis
of the object in order to plan a grasping movement is
different from and independent of that of the complementary reaching movement [7, 15]. In spite of this, a
similar mechanism of visual elaboration can be employed
to code the object of a grasping movement. That is,
the object can be encoded in an object-centred frame of
reference and successively transposed in an egocentric
one. In order to test this possibility, we devised an experimental paradigm similar to that used previously [8], in
which subjects were required to grasp a three-dimensional
bar superimposed over the shaft of the Maller-Lyer
illusion. We predicted that, if only a sequential analysis

of portions of the object coded in an egocentric frame of

reference is performed in order to select the opposition
points for the fingers, the grasping kinematic parameters
should not be affected by the illusion. On the contrary, if
at first a global analysis of the object is carried out, the
illusion effect should affect the grasping movement. A
successive transposition of the object in an egocentric
frame of reference should reduce the effect of the illusion.
Two control tasks were carried out in order to check the
perceptual effectiveness of the illusion configurations.

Methods
Eight right-handed subjects (two males and six females: age
20-31 years) participated in the present study. All were naive as
to the purpose of the experiment.
The subjects sat in front of a table with their right hand
resting on its surface. The starting position was a disc located
on the plane of the table along the subject's sagittal plane at
25 cm from his/her frontal plane. According to the task (see
below), at the beginning of each trial the subjects placed on the
starting position either the point of a pencil held in their right
hand (drawing task), or their thumb and index finger, held in
pinch position (grasping and matching tasks). Stimuli were
displayed on an easel located on the plane of the table, 35 cm
from the subject's frontal plane and inclined by 4 5 with respect
to the plane of the table.
Stimuli were the open and closed configurations of the
M~iller-Lyer illusion (Fig. 1). A non-illusory configuration in
which horizontal lines replaced either the open or the closed
wings was used as a control condition. Lines were 1 cm wide
and were drawn in black ink on the centre of white panels. The
shaft of the configuration lay approximately on the subject's
sagittal plane and could be either 5, 6 or 7 cm long. The ratio
between wings and shaft length was 0.3. A black wooden bar

Fig. 1. Experimental set-up. The upper and lower rows show the apparatus for presentation of the closed and open configurations,
respectively.

E. Daprati and M. Gentilucci/Grasping an illusion

of length equal to the shaft, and height of 1 cm, was superimposed over each shaft and fixed to the easel by means of a
magnet. Only the shaft was a three-dimensional object. Wings
were two-dimensional figures in order to avoid them becoming
an obstacle for the fingers during grasping movements. The
distance of the centre of the shaft from the starting position
was approximately 31 cm.
Three experimental sessions were run, separated by intervals
of at least I week. The tasks run in each session and the order
of their presentation were as follows.

Session 1: drawing task

Subjects were required to draw a vertical line of the same
length as the three-dimensional bar presented on the easel.
Inspection of the panel lasted for 5 sec, then the go signal was
given by a sound controlled by a PC. Vision of the stimulus
was allowed while drawing. On the contrary, vision of the hand
and the sheet on which the line was drawn was prevented by a
box covering them both.

Session 2. grasping task

Subjects were required to reach and grasp the extremities of
the three-dimensional bar presented on the easel with their right
thumb and index finger (Fig. 1). The procedure was the same
as in the drawing task, but full vision of both the stimulus and
the hand was allowed. The subjects were instructed to move as
naturally as possible, with the same velocity as during spontaneous movements. Removal of the bar from the easel was
easy because the strength of the magnet was weak.

1579

the subject's sagittal plane 8 cm from the edge of the table and
was used as a reference point.
In the grasping task, the time course of the distance between
the two markers positioned on the thumb and index finger was
used in order to study the grasping component of the movement. Maximal finger aperture, time to maximal finger aperture,
peak velocity of finger aperture and grasp time were the analysed parameters. The marker placed on the wrist was used to
study the transport component. Transport time, peak velocity
and deceleration time (expressed as percentage of total movement time) were measured. The distance between the markers
positioned on the thumb and the index finger was used in
order to measure finger aperture in the matching task. In both
grasping and matching tasks, the distance between the two
markers at the starting position was subtracted from that recorded at maximal finger aperture in the grasping task and at
finger aperture in the matching task in order to obtain the
distance between the finger palmar surfaces.
All parameters measured in the three tasks were submitted to
separate analyses of variance (ANOVAs) whose within-subject
factors were bar length (5 cm vs 6 cm vs 7 cm) and configuration
(open vs closed vs control configuration). We evaluated the
task's effect on the visual analysis of the bar length by comparing the illusion effect on the following parameters: drawn
line length (drawing task), maximal finger aperture (grasping
task) and finger aperture (matching task). We measured variation of these parameters for presentation of the open and
closed configurations with respect to the control configuration.
These values were submitted to an ANOVA, whose withinsubject factors were task (drawing vs matching vs grasping),
bar length (5 cm vs 6 cm vs 7 cm) and configuration (open vs
closed configuration). The Newman Keuls test was used as
post-hoe test in all analyses.

Results

Session 3: matching task

Drawin 9 task
Subjects were required to match the size of the three-dimensional bar presented on the easel by opening their right thumb
and index finger. The subjects were allowed to lift their hand,
but not to approach the object during the trial. The procedure
was the same as in the grasping task.
The aim of the first session was to establish whether the
modified configurations of the Mt~ller-Lyer figures would be
effective in inducing an illusion. The long interval between two
successive sessions was chosen in order to minimize practice
effects. Presentation order of the second and third task was
selected in order to prevent the perceptual judgement performed
in the matching task from influencing visuo-motor integration
in the grasping task.
In all tasks, the subjects were required not to pay attention
to the two-dimensional drawing upon which the bar was superimposed. In each session, 84 trials were run. That is, seven trials
for each stimulus length (5, 6, 7 cm) and configuration (open,
closed, control) were presented in pseudo-random order.
In the drawing task, the length of the drawn lines was manually measured by the experimenters at the end of the session.
Resolution was approximately 2 mm. Finger aperture in matching the bar length and the kinematics of reaching~rasping
movements were recorded using the ELITE system (B.T.S.,
Milan, Italy). Apparatus, movement reconstruction and data
elaboration procedures are described elsewhere [10]. The spatial
resolution of the ELITE system is 0.4mm [10]. Four markers
were used. The first marker was placed on the styloid process
of the radius at the wrist. The second and third markers were
positioned on the nails of the thumb and the index finger,
respectively. The fourth marker was placed on the table along

D r a w n lines were straight, and varied significantly

a c c o r d i n g to the three b a r lengths [F(2,14)=142.28,
P < 0 . 0 0 0 0 1 ; 31 . S m m vs 3 8 . 0 m m vs 45.0 mm]. H o w e v e r ,
subjects always u n d e r e s t i m a t e d the length o f the b a r (Fig.
2, u p p e r panel). D r a w n lines were clearly affected by the
illusion effect [F(2,14) = 9.96, P < 0.002]. T h a t is, a longer
line was d r a w n when the open c o n f i g u r a t i o n (39.4 mm)
was presented with respect to the c o n t r o l (38.0 m m ) a n d
the closed configurations (37.0 mm) ( P < 0.05).

Grasping task
Subjects o p e n e d their fingers progressively to a maxim a l finger a p e r t u r e a n d then closed on the object (Fig. 3).
Maximalfinger aperture was correctly scaled a c c o r d i n g to
the length o f the b a r to be g r a s p e d [57.6 m m vs 64.7 m m vs
72.9 mm; F(2,14) = 201.20, P < 0.00001 ]. This p a r a m e t e r
was significantly influenced by the illusion effect
[ F ( 2 , 1 4 ) = 4 . 8 1 , P < 0 . 0 5 ] , a l t h o u g h the effect was small.
Subjects o p e n e d their fingers wider when the open
(65.5 m m ) a n d the c o n t r o l (65.2 m m ) configurations were
p r e s e n t e d with respect to the closed c o n f i g u r a t i o n
( 6 4 . 5 m m ) (Fig. 2, lower panel, P < 0 . 0 5 ) . Peak velocity

E. Daprati and M. Gentilucci/Grasping an illusion

1580
D r a w i n g Task

Stimulus Length: 5 cm

65

100,
80 . . . . . .

g
c~O 45
,-d

,.

6o
40

....

0~ J
0

CL5 C05 OP5

. . . . . . .

2O

,-a

25

:!'

CL6 C06 OP6

CL7 C07 OP7

200

400
600
ms
Stimulus length: 6 cm

80

100,
Grasping Task

80

~'~ 85

60
~

40

.""

20
<

65

45

......... .

,,i"
0

200

400
600
ms
Stimulus length: 7 cm

100,
CL5 C O 5 0 P 5

CL6 C O 6 0 P 6

CL7 C O 7 0 P 7

80
Matching Task

....

60

85

,'"

40

g
-~ 65
<

45

800

CL5 C05 OP5

ill

I//

CL6 C06 OP6

CL7 C07 OP7

Fig. 2. Mean values of line length in the drawing task (upper

panel), of maximal finger aperture in the grasping task (middle
panel) and of finger aperture in the matching task (lower panel).
CL5=shaft length 5cm, closed configuration; CO5=shaft
length 5 cm, control condition; OP5 = shaft length 5 cm, open
configuration; CL6=shaft length 6cm, closed configuration;
CO6=shaft length 6cm, control configuration; OP6=shaft
length 6 cm, open configuration; CL7 = shaft length 7 cm, closed
configuration; CO7=shaft length 7cm, control condition;
OP7 = shaft length 7 cm, open configuration ( I = S.E.).

offinyer aperture, like maximal finger aperture, was greater for presentation of the open (258.4 mm/sec) and the
control (257.5 mm/sec) configurations with respect to the
closed configuration (248.8 mm/sec). However, this effect
did not reach significance. The remaining kinematic grasp
and transport parameters were not influenced by the
illusion.

20
0

i"~?

::
51'
0

200

400

600

800

ms

Fig. 3. Representative examples of time course of grasping the

extremities of the closed configuration (dotted lines), of the
control configuration (continuous lines) and of the open configuration (dashed lines) (subject CM.). One trial for each of
the three shaft length conditions is represented.

Across-tasks comparisons
As expected, a significant effect of the illusion was
found [F(1,7)=18.12, P<0.005; closed configuration
1.4 ram, open configuration 0.9 ram]. In addition, there
was a significant interaction between task and configuration [F(2,24) = 3.79, P < 0.05]. Post-hoc comparison
showed that the effect of the closed configuration was
significantly different in the matching task ( - 2 . 5 r a m )
with respect to the two other tasks (drawing task
- 1 . 0 r a m , grasping task - 0 . 7 m m ; P<0.05). No significant difference was observed for the open configuration among the three tasks (drawing task 1.4 mm,
grasping task 0.2 mm, matching task 1.1 ram).

Discussion

Matching task
Finger aperture was approximately scaled according to
the presented bar length [52.7 mm vs 61.7 mm vs 77.0 ram;
F(2,14)=36.79, P<0.00001]. Moreover, it was affected
by the illusion [F(2,14) = 6.85, P < 0.008]. Finger aperture
was wider for presentation of the open (65.4mm) and
the control (64.3 mm) configurations with respect to the
closed configuration (61.7ram) (Fig. 2, middle panel,
P<0.05).

The effect of the M/iller-Lyer illusion observed in both

the drawing and the matching tasks was smaller than that
observed in previous experiments in which perceptual
judgement on the length of the shafts of the two configurations was required [5, l l, 20, 21]. In the present
experiment, the three-dimensional shafts had large width
and therefore they did not produce blurring effects at
the vertices of the configurations [3]. Thus, the optical
component of the illusion was removed and only effects

E. Daprati and M. Gentilucci/Grasping an illusion

arising from visual analysis of figure shape could be elicited. It is known that we commonly use shape as an
indicator of size. In the drawing task, the length of all
figures was underestimated. This finding is likely a consequence of the lack of visual control while drawing. In
order to compensate for this, it is possible that the length
of the drawn line was judged on the basis of the time
elapsed since movement began. Since it is known that
hand movements without visual control of the hand are
performed slowly [10, 16], a possible slowing down of the
drawing could produce underestimation of the covered
distance.
The MMler-Lyer illusion influenced maximal finger
aperture during grasping. Like finger aperture in the matching task, this parameter was enlarged for presentation
of the open and control configurations with respect to
the closed configuration. The particular orientation of
the wings in the open configuration condition could be
an obstacle for the grasping fingers, thus accounting for
an increase in maximal finger aperture. However, since
only the shaft was a three-dimensional object, whereas
the wings of each configuration were two-dimensional
figures, we can exclude this possibility.
An effect of visual illusions on grasping movements
was also found in a previous experiment [1]. In that
experiment the two 'Titchener circle' configurations were
presented simultaneously and subjects were required to
pick up one of the two central discs according to a samedifferent judgement on their relative size. Such a task
required that the subjects visually analysed the scene
around the two potential target objects. This explains the
small effect of the illusion found by the authors. However,
due to the instruction given to the subjects, and to the
presentation of two configurations at a time, this experiment does not prove whether an analysis of the entire
target object and of the surrounding cues occurs for natural grasping movements.
The aim of the present experiment was to determine
the nature of the visual analysis that is performed in order
to grasp an object. We considered two possibilities. The
first possibility suggests that a serial analysis of adjacent
portions of the object is performed in order to select a
stable grasp. That is, the object is not analysed as a
whole, but it is fragmented in order to select successive
opposition axes, each computed with respect to the
observer (egocentric frame of reference). The second
possibility suggests that a global visual analysis, that
takes into account cues surrounding the target, is first
performed on the object. The findings of the present
experiment support this second hypothesis. Indeed, not
only was the distance between the two extremities of
the three-dimensional shaft of the MMler-Lyer figures
analysed, as suggested by the first hypothesis, but also
the remaining figure (the two wings) responsible for the
illusion. Interestingly, the configuration was considered
as a whole, although part of it was presented as a twodimensional figure. In other words, a process of visual
grouping, by which two-dimensional images were corn-

1581

bined with a three-dimensional object, was used in the

visual analysis of the target. This was true even for the
motor response.
The second hypothesis may also be supported by kinematic data collected on a visual agnosic patient and a
healthy control subject, picking up objects of different
shapes [18]. In this study the finger opposition axes, recorded on the object at the end of the grasp, frequently
passed through its centre of mass. The centre of mass of
an object can be established only if the whole object is
visually analysed. However, the opposite portions of the
object's surface corresponding to the selected finger
opposition axes were the only ones that were parallel to
each other, allowing a stable contact of the fingers on the
object. That is, these data may also support the first
hypothesis, which suggests an analysis of opposite surface
portions on the object.
In the present study the effect of the illusion was smaller
in the motor than in the reproduction tasks. This effect
was clear when the grasping task was compared with the
matching task, and less evident in the comparison with
the drawing task. This latter result may be due to the
more complex motor response required by the drawing
task, since both proximal and distal arm segments are
involved in drawing a figure. Such a complex motor
response might require a more efficient use of the egocentric frame of reference. This, in turn, might induce
greater correction of the illusion (see below). Another
possibility is that the effect of the illusion may have been
masked by the large underestimation of reproduction
amplitudes.
Taken together, these data suggest that, when visuomotor integration is performed, the effect of the illusion
is partially corrected. In a previous experiment we found
that, for pointing movements, the illusion effect is
inversely correlated with the efficiency of the egocentric
frame of reference [8]. We speculated that the effect of
the illusion, due to coding the target in allocentric coordinates, was partially reduced by successive transformation of visual information into motor commands,
that is while re-coding the object in egocentric coordinates. Although the visual analysis of the target has
been hypothesized as different and independent for reaching and grasping movements [7, 15], the results of the
present experiment suggest the possibility of there also
being two stages in object visual analysis for planning
grasp movements. We propose that in a first stage an
object is coded in an object-centred frame of reference,
whereas in a second stage it is transposed in an egocentric
one. In a previous neuropsychological study [9], we suggested that the parietal lobe is involved in transposing
the spatial position of the target of reaching movement
from an allocentric to an egocentric frame of reference.
On the basis of neurophysiological results [23], we can
suggest that an analogous transformation for grasping
movements is performed in the same parietal lobe. From
the AlP (anterior intraparietal area), an area involved in
planning grasp movements, neurons were recorded that

1582

E. Daprati and M. Gentilucci/Grasping an illusion

were activated by object fixation without grasping it

('object' neurons). In addition, other neurons, called
'non-object' neurons, were activated by vision of interaction of the hand with the object. One can speculate that
transformation of object coding from an object-centred
frame of reference (likely used to code an object by
'object' neurons) to an egocentric frame of reference (used
to code an object by 'non-object' neurons) can be performed by concurrent activity of the two types of
neurons.

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339-354.
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12. Iberall, T., Bingham, G. and Arbib, M. A., Opposition space as a structuring concept for the analysis
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Acknowledgements--We thank Dr P. Dominey for valuable

comments on the manuscript, and Drs M. Gangitano, G. Pagnoni and I. Toni for discussion of data. This work was supported by a Research Grant from the Human Frontier Science
Program, by a grant from the European Neuroscience Program
and by grants CNR (Centro Nazionale delle Ricerche) to the
Institute of Human Physiology of Parma and MURST (Ministero dell'Universit/t e della Ricerca Scientifica e Tecnologica)
to the Institute of Human Physiology of Parma and to M.G.

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