BR A IN RE S EA RCH 1 0 80 ( 20 0 6 ) 9 –1 6

a v a i l a b l e a t w w w. s c i e n c e d i r e c t . c o m

w w w. e l s e v i e r. c o m / l o c a t e / b r a i n r e s

Research Report

Dissociation of perception and action unmasked

by the hollow-face illusion

Grzegorz Króliczak a , Priscilla Heard b , Melvyn A. Goodale a,⁎, Richard L. Gregory c

a
Department of Psychology, Neuroscience Program, University of Western Ontario, London ON, Canada N6A 5C2
b
School of Psychology, University of the West of England, Bristol, UK
c
Department of Experimental Psychology, University of Bristol, UK

A R T I C LE I N FO AB S T R A C T

Article history: It has been suggested that there are two separate visual streams in the human cerebral
Accepted 10 January 2005 cortex: a ventral pathway that provides perceptual representations of the world and serves
Available online 6 March 2006 as a platform for cognitive operations, and a dorsal pathway that transforms visual
information for the control of motor acts. Evidence for this distinction comes from
Keywords: neuropsychology, neuroimaging, and neurophysiology. There is also evidence from
Hollow-face illusion experimental psychology, with normal observers experiencing an illusion—where
Vision-for-perception perception and action can be dissociated, although much of this evidence is controversial.
Vision-for-action Here, we report an experiment aimed at demonstrating a large dissociation between
Perceptual judgment perception and fast action using the hollow-face illusion, in which a hollow mask looks like
Visuomotor control a normal convex face. Participants estimated the positions of small targets placed on the
actually hollow but apparently normal face and used their fingers to ‘flick’ the targets off.
Despite the presence of a compelling illusion of a normal face, the flicking movements were
directed at the real, not the illusory locations of the targets. These results show that the
same visual stimulus can have completely opposite effects on conscious perception and
visual control of fast action.
© 2006 Elsevier B.V. All rights reserved.

1. Introduction The two-streams hypothesis has also received support

from somewhat controversial behavioral studies that have
The notion of an ancient visual system for rapid action and examined the influence of perceptual illusions on the control
a more recent visual system for (conscious) cognitive of object-directed actions such as saccades, reaching move-
operations, such as planning, makes evolutionary sense ments, and grasping (for reviews, see (Goodale and Milner,
(Goodale and Milner, 1992; Milner and Goodale, 1995). 2004; Goodale and Westwood, 2004)). Early experiments
Present evidence for these two streams–believed to be showed that saccadic eye movements are insensitive to a
organized into dorsal and ventral cortical pathways respec- dot-in-frame illusion in which the perception of a target's
tively–comes from rare patients with selective brain lesions location is shifted opposite to the displacement of a large
(Goodale et al., 1991; Perenin and Vighetto, 1988), neuroima- visual frame (Bridgeman et al., 1981; Wong and Mack, 1981),
ging studies (Culham and Kanwisher, 2001; Grill-Spector, suggesting that location is processed differently by the
2003), and neurophysiology (Cohen and Andersen, 2002; visuomotor and perceptual systems. Aglioti et al., (1995) later
Tanaka, 2003). demonstrated that the maximal opening of a grasping hand is

⁎ Corresponding author. Fax: +1 519 661 3961.

E-mail address: [email protected] (M.A. Goodale).

0006-8993/$ – see front matter © 2006 Elsevier B.V. All rights reserved.
doi:10.1016/j.brainres.2005.01.107
10 B RA IN RE S EA RCH 1 08 0 (2 0 0 6 ) 9 –1 6

insensitive to the robust perceptual illusion that a target disk by the dorsal system) would be directed to touch the truly
surrounded by smaller circles is larger than the same disk concave face, despite the presence of a compelling illusion of a
surrounded by larger circles (Ebbinghaus Illusion)—despite convex face (mediated by the ventral stream). To test this, we
the fact that grip opening is exquisitely sensitive to real asked participants to reach out rapidly and flick off targets
changes in the size of the target disk. Peak grasping aperture is that were placed on the mask.
refractory to this size contrast illusion even when the hand We also asked our participants to make slower and more
and target are occluded during the action (Haffenden and deliberate pointing movements to these same targets. We
Goodale, 1998), indicating that on-line visual feedback during did this because an earlier study had reported that pointing
grasping is not required to ‘correct’ an initial perceptual bias movements to a stereoscopic pair of pictures of a 3-D virtual-
induced by the illusion. reality hollow mask were directed at the perceived position
A number of recent findings, however, have challenged of the display, corresponding to the apparent reversed depth
the notion that perceptual illusions do not affect the control (Hartung et al., 2005). There is evidence, however, that
of object-directed actions. These challenges fall into several pointing movements can often be influenced by cognitive
categories including: non-replication (Franz et al., 2003), the factors and need not engage the ‘automatic’ mechanisms in
contention that early studies did not adequately match the dorsal stream (Bridgeman et al., 1997). In fact, neurolog-
action and perception tasks for various input, attention, ical patients with dorsal-stream damage, who cannot reach
and output demands (Bruno, 2001; Smeets and Brenner, 2001; out accurately to acquire targets using rapid ‘automatic’
Vishton, 1999), or the idea that action tasks involve multiple movements, can sometimes improve their performance
stages of processing from purely perceptual to more ‘auto- dramatically if they are encouraged to slow down and
matic’ visuomotor control (Glover, 2004; Glover and Dixon, move more deliberately, thereby (it has been argued)
2001). Some of the competing accounts (Glover, 2004; Smeets engaging ‘perceptual’ mechanisms in the ventral stream
and Brenner, 2001) are difficult to separate from the original (Rossetti et al., 2005). This might explain why the partici-
two-streams proposal. In addition, some of the contradictory pants in the earlier study Hartung et al., (2005) pointed to the
findings (Glover and Dixon, 2001) can be explained by perceived (i.e., the illusory) location of the mask. We
appealing to the fact that illusions can arise at different expected our participants to behave the same way when
stages in visual processing (Dyde and Milner, 2002). Accord- making slow and deliberate pointing movements to the
ing to this argument, illusions that arise in early visual areas, targets placed on the mask. In contrast, we expected that
such as primary visual cortex, will have an effect on action, participants would direct their rapid flicking movements to
whereas illusions that arise at higher stages of visual the veridical locations of the targets, showing no sensitivity
processing in the ventral stream will not. Nevertheless, to the concurrent illusion of depth.
because the illusory distortions that have been used in all
the studies to date were no more than a few millimeters, the
experiments were technically difficult to carry out and this 2. Materials and methods
could explain why the results have sometimes been hard to
replicate (Franz et al., 2003). Thus, a large visual illusion 2.1. Participants
(preferably many centimeters) that nevertheless shows a
clear dissociation between perceptual report and action Eight right-handed participants (mean age 23 years) were
would be reassuring. tested. All participants reported normal or corrected-to-
We looked for a dissociation between conscious percep- normal visual acuity and were either compensated financially
tion and rapid action using the large and dramatic depth for their time or given a course credit. They gave their
reversal of the hollow face, in which a realistic hollow informed consent prior to testing. The experiment was
mask appears as a convex face (Gregory, 1970). This is approved by The Review Board for Non-Medical Research
evidently a knowledge-based, top–down effect, where Involving Human Subjects (at the University of Western
extensive and powerful (though implicit) knowledge of Ontario) and was carried out in accordance with the principles
convex faces rejects the correct hollow perception in favor of the Helsinki 1964 Declaration.
of reversed depth (Gregory, 1997). According to the two-
streams hypothesis, the cognitive hollow-face illusion 2.2. The experimental conditions and responses
arises within the ventral stream and should therefore not
affect visuomotor computations in the dorsal stream (Dyde There were three experimental conditions: (1) a normal
and Milner, 2002). Although the hollow-face illusion is convex face mask seen as a normal face, (2) a hollow mask
strong, it can be countered by powerful opposed bottom–up seen as an illusion—looking convex and not hollow, and (3) a
information, especially binocular information from close hollow mask seen as a hollow face.
viewing with both eyes (Hill and Bruce, 1993), or less There were three measured responses: (1) fast “flicking”
effectively by shape-from-shading, with strong overhead movements with the finger to targets on the face, (2) slow
illumination of the hollow mask (Hill and Bruce, 1993; pointing movements with the finger to the perceived locations
Ramachandran, 1988). of targets on the face, and (3) drawing of the target positions
The main question was this: would rapid hand movements on paper (relative to a reference plate). In all three cases, the
be directed to touch the real position of the mask or (wrongly) dependent measure was the horizontal displacement (dis-
to the illusorily reversed mask? The two-streams hypothesis tance in the Z dimension of depth) with respect to the
would predict that fast (and automatic) movements (mediated reference plate.
 BR A IN RE S EA RCH 1 0 80 ( 20 0 6 ) 9 –1 6 11

2.3. The mask displays and their presentation optimal in three individuals tested in an earlier pilot study.
Subsequently, stronger or weaker lenses were employed until
The stimuli were two female face masks, identical except that the participant reported a strong and immediate illusion. As it
one was convex and the other concave (22.8 cm long and 14.8 turned out, the mean lens strength used in the experiment
cm wide). Sheet metal was glued to the backs of the masks, so was also −3.75 diopters.
small target magnets could be placed on their front surfaces. Testing took place in a dark room where the only source of
The faces were mounted on a ‘reference plate’, such that the illumination was a hidden spotlight. The direction of lighting,
normal (convex) face protruded in front and the hollow face and/or its intensity, was different for each experimental
receded behind it. The displays were mounted firmly on a condition. The hollow mask, seen illusorily as convex, was
rotatable turret as shown in Fig. 1. The device allowed us to illuminated from below by a small spotlight. To keep shadow
present one mask at a time by rotating it to either one of two information similar, the spotlight was placed above the
settings. normal mask. In one final condition, the illusion for the
A sliding mechanism allowed us to move the entire display hollow mask was abolished by bright overhead lighting and by
on each trial to one of 3 randomized distances (19.8, 24.8, and removing the de-focusing lens.
29.8 cm from the start button) to prevent participants from
making stereotyped movements. A single target–a small 2.5. Procedure
cylindrical magnet (0.4 cm long and 0.5 cm in diameter)
covered with white cloth tape–was presented at two different Each behavioral measure (fast flicking, deliberate pointing,
depth locations on the faces: the side of the cheek (1.1 cm from and paper-and-pencil drawing) was tested in separate blocks
the reference plate) or the forehead (5.5 cm from the reference of trials. In the first part of the experiment, the normal face
plate). The displays were sufficiently large that participants and hollow face looking illusory were randomly interleaved in
could flick the targets from the hollow face without colliding a different order for each participant. In the last part of the
with the edge of the mask. In other words, the same type of experiment, the hollow face looking hollow was presented on
reaching movements could be used to flick targets off either its own.
the normal or the hollow face. The fast flicking and slow pointing were performed in
visual open loop (no visual feedback after the finger left the
2.4. The viewing conditions start button). LCD (PLATO) goggles were used to control the
viewing time: the face display was revealed and then, after 3 s,
The hollow-face illusory depth reversal is robust, except when a start signal was given to initiate the participant's fast
countered by strong stereo information in near viewing with flicking, or slow pointing, movement to the target. The goggles
both eyes. As the mask had to be sufficiently near in this became opaque as soon as the moving finger left the start
experiment for the participant to reach the targets, this was a button.
problem. Hence, we reduced the countering stereo by filtering Fast flicking: Participants were asked to flick the small
out high spatial frequencies, with a de-focusing lens placed magnet off the face as quickly and accurately as they could,
over the non-dominant eye, which allowed only low-frequen- using their index finger. These flicking movements were
cy binocular information. The lens was selected individually measured with an optoelectronic system (OPTOTRAK 3020:
to preserve the illusion within reaching distance. The first lens Northern Digital), which recorded (at 200 Hz) the position of an
tried was always −3.75 diopters, a value that was found to be infrared emitting diode located at the base of the index finger.

Fig. 1 – Left panel. The apparatus used to present the normal and hollow faces. Two small magnets, which served as targets,
are shown in position on the forehead and cheek of the normal face. These targets were always presented separately. Right
panel. The front view of the hollow mask. The lighting for this face comes from below, creating a shadow pattern that is similar
to lighting from above for the normal faces. The reader should see this hollow mask as an illusory convex face.
12 B RA IN RE S EA RCH 1 08 0 (2 0 0 6 ) 9 –1 6

In the first part of the experiment, there were 96 trials, 48

towards the illusory and 48 towards the normal face, 24
towards the cheek, and 24 towards the forehead. That is, there
were 8 trials at each of the three distances for each target
position. Participants were given 6 practice trials before
beginning the experiment, 3 with the hollow (illusory) face
and 3 with the normal face.
Deliberate pointing: Participants were instructed to point
directly to the location where they perceived the target. On
other trials, they were instructed to point the same
corresponding distance below the face (to avoid the possibility
of tactile feedback, particularly in the case of the normal face).
These slow pointing movements were also recorded with the
OPTOTRAK. In the first part of the experiment, there were 96
trials, 48 towards the display, and 48 below the display. The
order of the testing conditions was randomized across
participants. Within each condition, both viewing distance Fig. 2 – Perceptual estimates. Participants drew on a sheet of
and face display were also randomized, with a different trial paper the perceived positions of the target on the foreheads
order for each participant. Because the responses made and cheeks of the faces. The mean distances of the pencil
towards and below the display did not differ from one another, marks (along the horizontal z axis) from the line indicating
the results from both conditions were combined in subse- the reference plate are shown on the ordinate. Error bars
quent analyses. indicate standard errors of the mean.
Paper-and-pencil drawing: After viewing the face for 3 s,
participants drew the perceived position of the target on a looking hollow were veridical, i.e., reversed with respect to the
piece of letter-sized paper by placing a mark to the right (for normal face (Fisher–Hayter, P < 0.01).
near) or left (for distant) of a vertical reference line represent-
ing the reference plate of the mask. A new piece of paper was 3.2. Fast flicking
presented on each trial, and participants drew the position of
only one target on that piece of paper. In the first part of the Participants were equally fast at initiating their flicking move-
experiment, participants were tested 24 times for each target ments to the illusory (504 ms, SE = 32 ms) and the normal face
position (cheek and forehead), i.e., 8 times at each of the three (507 ms, SE = 34 ms), suggesting that the presence of the
distances for the convex and concave mask in a different illusion did not slow down the programming of the required
random order for each participant. movements. The movement onset time with the hollow face
The perceptual measures of the perceived target distances looking hollow was significantly faster (448 ms, SE = 31 ms;
(both drawing and deliberate pointing) were each performed F(2,14) = 8.3, P < 0.01; Fisher–Hayter, P < 0.01), an improvement
in two separate blocks, half of the trials before and half after which probably reflects the absence of the de-focusing lens
testing the fast flicking movement. This was done to test and the brighter viewing conditions on trials in which the
whether or not performing the visuomotor task would hollow mask was seen as hollow (Jiang et al., 1991).
influence the way the target positions (and as a result, the As Fig. 3 shows, in all three cases, the movements were
illusory display) are perceived. As it turned out, this made no directed to the real position of the targets on the cheek and
difference, so the results were collapsed. forehead. Thus, the end points of the flicking movements to
In a final set of trial blocks, participants were tested on all both the illusory (hollow) face and the hollow face looking
three tasks with the hollow face looking hollow. Since only hollow did not differ from each other but both differed
one type of display (the hollow mask) was presented, the significantly from the end points of the movements to the
number of trials was halved. In all other respects, the testing normal face (F(2,14) = 139.6, P < 0.001).
was identical. Indeed, the distance at which the target was flicked from
the forehead of the normal face was more than 9 cm closer to
the participants than the distance at which it was flicked from
3. Results the forehead of either the hollow face looking hollow or the
hollow face looking normal (the illusion). The flicking on both
3.1. Drawing the normal face and the hollow face looking hollow corre-
sponded to the seen positions within about a centimeter; but
As can be seen in Fig. 2, when participants drew the position of for the hollow face looking normal (illusion), the flicking for
the target presented on the forehead or cheek of the different the forehead was about 7 cm behind its seen position. In other
masks, they showed evidence of a robust hollow-face illusion words, for the hollow-face illusion, the perceived position of
(F(2,14) = 57.4, P < 0.001). In other words, the relative positions the target went one way and the final position of the hand in
of the target on the illusory display were the same as they the flicking task went the other. Finally, it should be noted that
were on the normal face mask, although the distance between the flicking responses to the hollow-face illusion showed no
them was seen as slightly compressed (Fisher–Hayter, P < evidence of improving over time (F(11,77) = 0.86; ns) and that
0.01). The relative positions of the targets on the hollow face participants failed to hit the target on about half the trials.
 BR A IN RE S EA RCH 1 0 80 ( 20 0 6 ) 9 –1 6 13

the hollow face appear hollow (Jiang et al., 1991). But even in
this condition, participants failed to hit the target on
approximately 30% of the trials.
Fig. 5, which shows the paths of the flicking movements
(seen from the side), also makes the point that participants
were programming their responses differently for the normal
and illusorily depth-reversed faces. Note that the trajectories
for these two conditions separated right from the start of the
movements. Indeed, the average trajectory for movements
made to the illusory face was much more similar to the
average trajectory for the hollow face looking hollow than it
was to the average trajectory for the normal face.

3.3. Slow pointing

There were clear differences in the movement onset times for

Fig. 3 – The flicking task: the mean distance of the hand (along pointing with the three different displays (F(2,14) = 7.4, P <
the z axis) at the moment the participant attempted to flick off 0.001). The mean onset time for pointing movements to the
the target from the cheek or forehead of the normal or hollow illusory (hollow) face (737 ms, SE =63 ms) was significantly
mask. Notice that, in the case of the illusory face, the end longer (Fisher–Hayter, P < 0.05) than the mean onset time for
points of the flicking movements corresponded to the actual movements to the normal face (692 ms, SE = 53 ms), and both
distances of the targets, not to consciously seen distances were significantly longer (Fisher–Hayter, P < 0.01) than the
(Fig. 2). Error bars indicate standard errors of the mean. onset time for pointing movements to the hollow face looking
hollow (612 ms, SE = 44 ms). Movement times did not differ
Fig. 4 shows that, early in the flicking movement, at the across the three conditions. The average duration of the
point of maximum velocity, the horizontal (z axis) distance pointing movements (1660 ms, SE = 155 ms), however, was
covered on the way to the target was greater for both the more than three times longer than the average duration of the
hollow face looking hollow and the hollow-face illusion than it flicking movements (471 ms; SE = 30 ms).
was for the normal face (F(2,14) = 37.7, P < 0.001 and Fisher– As Fig. 6 shows, the final positions of the pointing move-
Hayter, P < 0.01). ments made to the illusory (hollow) face, like those made to
It should be noted, however, that the distance reached on the normal face, were in front of the reference plate. In
trials with the hollow face looking hollow was slightly greater contrast, the final positions of the pointing movements made
than it was for the illusory face (Fisher–Hayter, P < 0.05), which to the hollow face looking hollow were located beyond the
again probably reflects the absence of the de-focusing lens and reference plate (F(2,14) = 203, P < 0.0001). The final positions of
the brighter viewing conditions that were required to make the pointing movements made to the illusory face were
somewhat closer to the reference plate as compared to the
final positions of movements made to the normal face (Fisher–
Hayter, P < 0.01) and also did not reflect the perceived relative

Fig. 4 – The mean distance (along the z axis) from the start
button reached at maximum velocity for the flicking Fig. 5 – A side view of the paths of movements in the flicking
movements made to targets placed on the three different task in the three conditions. The profiles show position of the
displays. When maximum velocity was reached, early in the moving finger in the y (vertical) and z (depth) axes. The IRED
movement trajectory, participants were already reaching out placed on the base of the index finger was tracked at 200 Hz
further for both the hollow faces, however they appeared, (and the obtained data were then normalized). The mean
than they were for the normal face. Error bars indicate distance of the reference plate of the display was 24.8 cm
standard errors of the mean. from the start button.
14 B RA IN RE S EA RCH 1 08 0 (2 0 0 6 ) 9 –1 6

other words, the motor system was not initially fooled by the
visual illusion, as participants did not move their hand to the
perceived position of the target first, correcting their move-
ment later. It is also apparent that the participants were not
moving their finger “blindly” forward until they made physical
contact with the mask. Instead, they appear to have
programmed the distance to be moved quite early on. As a
consequence, the trajectories for the two kinds of displays–
illusory and normal–separated right from the start. In other
words, it was not the perceived position of the target but its
actual distance that determined the movement trajectory.
The results from the drawing test show that the illusory
face was perceived as somewhat compressed. This effect,
which has been reported by others (Hartung et al., 2005), may
reflect the operation of Emmert's Law, in which the apparent
size of the protruding features such as the forehead would
Fig. 6 – The pointing task: the mean final location of the
appear smaller for the illusory than for the real face. In any
finger with respect to the reference plate for pointing
case, the fact that the illusory face appeared somewhat
movements made to the cheek and forehead targets for all
compressed and that there were other cues, such as reverse
three displays. Notice that the movements made to all three
motion parallax, meant that participants could have distin-
displays tended to end near the perceived positions of the
guished between the illusory and normal mask. In principle,
targets, although the movements to the targets on the illusory
then, the participants could have used this knowledge to make
face did not distinguish between the cheek and forehead
larger amplitude movements when presented with the
targets. Error bars indicate standard errors of the mean.
illusory face. But this seems unlikely. First, the position of
the display was randomly changed from trial to trial making it
positions of the forehead and cheek targets. Nevertheless, as difficult to use this strategy. Second, they took no longer to
Fig. 7 illustrates, the pointing movements were typically initiate their flicking movement to the targets on the illusory
directed to locations in front of the reference plate, in sharp display than they did to the targets on the normal face. In
contrast to the flicking movements, which were always addition, there was no evidence that participants learned to
directed at the real position of the targets deep inside the increase the amplitude of their flicking movements to the
mask (F(2,14) = 27.7, P < 0.001). illusory face as the experiment progressed. Finally, we have

3.4. Effect of flicking on perceptual estimates

Neither the drawing estimates nor the pointing movements

were affected by prior performance of the flicking task (F(1,7) =
0.3, ns; and F(1,7) = 1.6, ns, respectively). In other words, there
was no difference between the perceptual measures collected
before the flicking task and the measures collected afterwards.
This demonstrates that the haptic feedback from the veridical
position of the target on flicking trials did not influence
perception of the hollow-face illusion.

4. Discussion

The results of the experiment demonstrate that, despite the

presence of a strong hollow-face illusion, participants directed
rapid movements to the real, not the illusory positions of the
targets. To do this, the visuomotor system must either have Fig. 7 – The slow pointing and fast flicking responses (the
access to other sources of visual information or process the endpoints are averaged over both target positions). For the
available information differently from that driving the illu- normal face and the hollow face looking hollow, both the
sion. Participants seem to be unaware of the veridical depth flicking and slow hand movements were nearly veridical. For
information they are using to control their flicking move- the hollow face looking convex (the illusory reversal of
ments. Moreover, the use of this veridical depth information depth), the movements were very different. Here, the end
does not ‘break’ the illusion. points of the slow pointing corresponded to the illusory
The analysis of the trajectories of the finger during target- position of the target (in front of the reference plate) whereas
directed flicking movements shows that these movements the end points of the fast flicking movements corresponded
were ‘programmed’ using the real position of the target—even to their actual position (behind the reference plate). Error
on trials in which the hollow-face illusion was present. In bars indicate standard errors of the mean.
 BR A IN RE S EA RCH 1 0 80 ( 20 0 6 ) 9 –1 6 15

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