Perception, 2001, volume 30, pages 945 ^ 957

DOI:10.1068/p3088

Mental imagery of visual motion modifies the perception

of roll-vection stimulation
Fred W Mast, Alain Berthoz, Stephen M Kosslyn

Department of Psychology, Harvard University, William James Hall, 33 Kirkland Street, Cambridge,
MA 02138, USA; e-mail: fmast@wjh.harvard.edu; Laboratoire de la Physiologie de la Perception
et de l'Action, CNRS et College de France, 11 place Marcelin Berthelot, F 75005 Paris, France
Received 8 June 2000, in revised form 2 May 2001

Abstract. When viewing a wide-angle visual display, which rotates in the frontoparallel plane
around the line of sight, observers experience an illusory shift of the direction of gravity; this
shift leads to an apparent tilt of the body and displaces allocentric space coordinates. In this
study, subjects adjusted an indicator to the apparent horizontal while viewing a rotating display.
To determine whether top ^ down processes could affect the illusion, the subjects were asked to
visualize a rotating configuration of dots onto a blank central portion of the moving visual field.
Visualizing dots and actually viewing the dots deflected the spatial judgment in very similar ways.
These results demonstrate that top ^ down processing can affect allocentric space coordinates.

1 Introduction
Perception requires the brain to produce a coherent and unique percept on the basis
of ambiguous sensory information. Because the input often underspecifies the nature of
the stimulus, our perceptual systems sometimes give rise to illusions and other sorts of
errorsfor example, when sitting in a stationary train while watching the neighboring
train pull out of the station. This phenomenon is called visually induced self-motion, or
vection (Mach 1875; Fischer and Kornmuller 1930). This sort of linear vection (Young
et al 1973; Berthoz et al 1975; Pavard and Berthoz 1977) occurs because whole-scene
motion signals induce self-motion perception, and the sensory inputs arising from
steady motion of the self through a stationary environment are in fact identical to
those produced by a moving scene past a stationary observer.
In addition, rotating patterns produce, in the simplest case, circular vection, which
can be elicited when the surrounding pattern rotates around the mid-body axis of an
upright observer. In this case, observers feel as if they themselves are rotating in the
opposite direction to the actual rotation. Moreover, rotations of the scene around
the line of sight produce roll vection. Dichgans et al (1972) used a display of random
texture, which contained no orientation cues to the visual vertical to produce such
roll vection; this stimulation induces the visual sensation that the observer is continuously rotating, which conflicts with extraretinal gravity-receptive information signaling
no change in posture. This conflicting sensory information results in an illusory shift
of the internal representation of the direction of gravity, which leads observers to feel
that their bodies are tiltedand also leads observers to perceive a vertical line as tilted
in the same direction. Studies in weightlessness have shown even stronger effects of
roll-vection stimulation than occur when gravity is present (Young et al 1986a, 1986b).
Studies of vection generally emphasize the role of sensory stimulus properties, and
focus almost exclusively on bottom ^ up processing. For example, much has been
discovered about the role of velocity and spatial frequency (Lestienne et al 1977), area
and size of stimulation (Brandt et al 1973), relative motion and distance between
foreground and background (Howard and Heckmann 1989; Howard and Howard 1994;
Nakamura and Shimojo 1999), and the type of visual motion pattern used to elicit vection
(Andersen and Braunstein 1985). In addition, the neural mechanisms underlying these

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F W Mast, A Berthoz, S M Kosslyn

properties of vection have been studied in the monkey. For example, researchers have
shown that, during circular horizontal motion, visual motion information and vestibular
information converge on vestibular nuclei neurons, which respond to visual motion
information in one direction and to vestibular information arising from whole body
rotations in the opposite direction (Henn et al 1974). A wealth of information has
implicated brain stem mechanisms in self-motion perception. Furthermore, integrated
multisensory signals from the vestibular nuclei clearly project to areas of the parietal
lobes that are involved in spatial orientation (eg Grusser et al 1990). Neuroimaging
studies in humans have identified the human analog of the multisensory parieto-insular
vestibular cortex (Bottini et al 1994; Lobel et al 1998).
Based on such findings from neurophysiology and psychophysics, the dominant
theory of how visual motion information and vestibular information are integrated
focuses on bottom ^ up processing. To our knowledge, the possibility that cognitive factors
might influence space perception has been widely disregardedin spite of sporadic
suggestions such as those by Lackner and Levine (1979), who documented that perception of spatial orientation depends in part on the context. More recently, Mergner
et al (2000) found that specific instructions to the subjects created different perceptual
states during circular vection; specifically, in one condition, they asked their subjects
to `stare through' the optokinetic pattern, and, in another condition, they asked them to
accurately track the moving pattern with the eyes. They found that the latter condition
usually eliminated reports of vection. In the present study, we ask whether top ^ down
processing via visual mental imagery can affect the perception of ambiguous visual and
vestibular (or proprioceptive) signals during roll-vection stimulation. Top ^ down processing occurs when one's knowledge or beliefs affect relatively low-level processes.
The hypothesis that visual mental imagery can modulate the perception of rollvection stimulation is consistent with reports that imagery can interfere with low-level
visual processes, such as vernier acuity (Craver-Lemley and Reeves 1987). Indeed,
responses of individual neurons in area V1 can in fact be modulated via attention
(Motter 1993), and neuroimaging studies have shown that visual mental imagery can
activate this area (Kosslyn et al 1995, 1999). Neuroimaging techniques have further
revealed that top ^ down processing can affect motion processing. For example, in a
functional magnetic resonance imaging (fMRI) study, O'Craven et al (1997) found
a significant increase in activation in the human analog of the MT ^ MST complex
when the subjects were attending to moving dots compared to when they were attending
to stationary dots within the same visual stimulus.
However, none of the previous studies dealt with the possible role of visual mental
imagery in mediating top ^ down influences on motion processing. To our knowledge,
the present study is the first in which the effect of imagined motion on the perception
of allocentric orientation has been investigated. Allocentric orientation tasks require
the subjects to make spatial judgments with respect to an external reference axis. In a
previous study, we (Mast et al 1999) showed that, when subjects visualized gratings in
a particular orientation, the perceived gravitational vertical was deflected in a very
similar way as when the subjects actually viewed the gratings. These results, however,
were based solely on the use of stationary gratings in imagery and perception. In the
present study, subjects visualized moving dots rather than stationary gratings, and our
focus is on discovering whether imagery can affect vection per se.
The dearth of research on the role of top ^ down processing in the perception of
roll vection is surprising, given that most models of spatial orientation include some
sort of top ^ down component, such as a central processor or an internal model as
proposed by Merfeld et al (1999). But there is only very limited support for the existence
of such processing (Berthoz and Viaud-Delmon 1999). To test the role of imagined
motion in modulating motion perception, we used a paradigm similar to Dichgans

Mental imagery and vection

947

et al's (1972), but in which visual mental images of a moving pattern could interfere
with the integration of afferent sensory information.
2 Methods
2.1 Apparatus and stimuli
The subjects were tested individually. They sat upright, with their heads restrained in
a chin rest, facing a cylindrical drum that rotated around a horizontal axis. The inner
sides of the drum contained randomly positioned spots, which covered an annular
area between 54 and 80 deg of the subject's visual field. A black mask excluded all
potentially visible cues that were outside the wide-angled drum (visual angle: 80 deg).
The interior of the drum was illuminated from outside by diffuse tungsten light. The
subject's head remained restrained throughout the task, which also enhanced the experience of vection. The height of the chair was adjustable, so that the center of
the drum could be set at eye level for each subject. Each spot subtended a visual angle
of approximately 4.4 deg. The drum could be rotated at a constant velocity around
the line of sight in either the clockwise or counterclockwise direction. The angular
velocity of the optokinetic stimulation by the moving drum was adjustable and held
constant to 30 deg s1 in this study; a rotational velocity between 30 and 40 deg s1
was previously shown to have the maximal effect on the perception of visual orientation (Held et al 1975). Disks with different patterns could be mounted onto the central
portion of the drum (visual angle: 54 deg).
A first disk contained eighteen spots, arranged in two concentric circular bands,
with six spots in the inner band (with a visual angle of 18 deg) and twelve spots in the
outer band (with a visual angle of 38 deg), as illustrated in figure 1. A second disk
contained only six spots arranged in the inner band. A third disk contained only a
fixation spot in its center, in line with the drum's axis of rotation; an identical fixation
spot was also placed in the center of each other disk. We used these stimulus configurations because they can be easily grouped, memorized, and later visualized in the
imagery conditions. All the disks were presented in combination with the motion cues
in the far periphery as described above (randomly positioned spots). In particular,

counterclockwise

rotation of
moving pattern
clockwise

Figure 1. The pattern with both circles (3 and 4) was used for the complete-perception condition.
Each spot subtended a visual angle of 5.5 deg. Two small laser dots (2) were superimposed on
the disk and could be rotated independently on a circular path subtending 28 deg in diameter.
The fixation spot (1) was positioned right in the middle of the pattern. In a second perception
condition partial perceptionthe outer circle (4) was absent.

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F W Mast, A Berthoz, S M Kosslyn

this motion information served as a perceptual aid when subjects imagined the
patterns in motion. The size of these stimulus configurations was similar to those that
have previously been found to induce vection by Andersen and Braunstein (1985),
Howard and Heckmann (1989), and more recently by Allison et al (1999).
A small keypad with push buttons controlled a motor, which allowed subjects to
adjust a visual indicator, as illustrated in figure 1. The visual indicator consisted of
two laser dots, which were projected onto the central portion of the drum at equal
distances from the fixation spot. The subjects were instructed to adjust the two lasers
so that they would appear aligned with the gravitational horizontal. The frame that
held the laser pointers could also be controlled by the investigator. The laser pointers
were randomly displaced before they were presented again. The orientation of the
indicator was measured by a potentiometer (accuracy 0.18).
In an earlier study we showed that two light sources could be used to perform
this task (Mast et al 1999), which has the advantage of producing minimal interference
when subjects form mental images. The laser dots were the only stationary cues visible
to the subjects. The frame holding the laser pointers was placed immediately behind
their heads. By operating the two push buttons, the subjects could orient the two laser
dots on a circular path (visual angle: 28 deg). The adjustable range was 258 with
respect to the physical horizontal, which was sufficient to register the perceptual biases
induced by the rotating patterns; the limited range of motion occurred because of the
subject's head and torso, which partly occluded the projection of the laser pointers
from behind. Minor differences have been reported when subjects set the indicator to
the apparent horizontal compared to the apparent vertical (Mast and Jarchow 1996;
Betts and Curthoys 1998; Van Beuzekom and Van Gisbergen 2000), but they are not
relevant in the context of the present study, in which we focus on the effects induced
by perceived and imagined visual motion. After the subjects had adjusted the lasers
appropriately, they pressed a button on the keypad, at which point the laser dots were
shut off. The keypad could be disabled by means of a switch on the control unit,
which was operated by the investigator. During the experiments, there were no other
stationary visual cues that could interfere with the task.
2.2 Subjects
Eighteen people (nine female, nine male, mean age 22 years, range 18 ^ 28 years)
volunteered to take part in this study, which was conducted in William James Hall
on the Harvard University campus. The subjects gave their informed consent and
completed a health history questionnaire prior to taking part in this study. None of
them reported any health problems and all subjects had normal or corrected-to-normal
vision. The subjects were Harvard students or professionals from the Boston area.
All subjects were na| ve regarding the purpose of this study and they were paid for
their participation. The study was approved by the Harvard University Institutional
Review Board.
2.3 Procedure
2.3.1 Perceptual conditions. Before the disk started to move, the two laser points appeared
in a random orientation and the subjects set them so that they appeared aligned with
the gravitational horizontal. The stationary spot patterns provided no orientation cueing.
The instructions ensured that subjects understood the task (``Adjust the two dots so that
they appear horizontally aligned, that is, so that the two dots are equally distant from
the ground'' and ``If a small ball were placed on an imaginary line connecting the
two dots, it would rest still and neither roll toward the left nor toward the right dot'').
We refer to these adjustments as subjective visual horizontal (SVH).
First, when the drum started to rotate, we measured the latency of vection onset
by having the subjects press a button at the first sign of perceived body motion.

Mental imagery and vection

949

Then, during drum rotation at 30 deg s1 constant velocity, the subjects performed
eight adjustments of the SVH. After the rotation stopped, the testing room was fully
illuminated and the subjects rested for approximately 5 min in order to eliminate
possible motion aftereffects before the next condition began. The subjects viewed all
three different disks described above (see figure 1). The disks were presented in a counterbalanced order. The condition in which we presented the disk with the full set of
spots (arranged in two bands) will be referred to as the complete-perception condition;
and the condition in which we presented the disk with only one band of spots will be
referred to as the partial-perception condition. The condition, in which we presented the
disk with only the fixation spot in its center will be referred to as the surround-motion
condition because the only motion information is due to the randomly positioned spots
in the far periphery. The top row in figure 2 provides an overview of the perceptual
conditions. Finally, half the subjects viewed clockwise rotating patterns and half viewed
counterclockwise rotating patterns.
2.3.2 Imagery conditions. After completing the perceptual conditions, the subjects
were asked to perform two imagery tasks. Prior to each, they participated in a training
session during which they inspected the pattern with eighteen spots for 30 s.
One imagery condition will be called the image-generation condition. In this
condition, we asked the subjects to visualize the full pattern, with all the spots, as
accurately as possible while they viewed the black disk with only the fixation spot on it.
When they reported having generated the image as accurately as possible, they were
again shown the pattern and asked to correct their mental image if necessary. This
procedure was repeated five times; at the conclusion of this training session, the subjects
reported being able to visualize accurately the spots moving.
A second imagery condition will be referred to as the image-completion condition.
In this condition we presented the pattern with the six spots arranged in the inner
band and the fixation spot in the center. The subjects were instructed to visualize the
outer band of spots at the same time they were seeing the inner band of spots, as if
they were seeing the entire pattern. Hence, this condition differed from the imagegeneration condition in that there were more cues visible while the image was formed.
As in the image-generation condition, the image-completion condition began with a
learning phase during which the subjects visualized the pattern repeatedly and corrected
their mental image if necessary. The subjects kept their gaze centered on the fixation
spot when they visualized the stimuli.
In both imagery conditions, after the subjects formed the image the drum started
to rotate, and subjects practiced visualizing the pattern in motion, as if it was painted
on the rotating drum. This procedure was repeated until they could maintain the
visualized motion pattern for at least 20 s. The velocity of the drum was held constant
at 30 deg s1. In both imagery conditions, just as in the perceptual conditions, the
subjects indicated vection onset before they performed the adjustments of the SVH.
The subjects required between 5 and 10 s to adjust the laser dots, indicating the SVH.
When the laser pointers were off, the subjects kept on looking at the fixation spot
but they were not required to maintain the mental image.
The two imagery conditions were presented in a counterbalanced order over subjects. However, the imagery conditions were always performed after the perception
conditions, which also provided an additional opportunity to learn the locations of the
spots. In pilot studies, we explored the possibility of having the imagery conditions
precede the perception conditions, but the subjects reported that the extended imagery
training sessions biased their settings in a subsequent perception condition, in which
the spots were absent. They claimed that it was difficult not to visualize after being
trained to form the mental images repeatedly. Therefore, we decided to keep the order

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F W Mast, A Berthoz, S M Kosslyn

constant, and the perception conditions were always completed first. That said, we
were concerned that this design feature might produce a possible carry-over effect
from previously setting the indicator in the perception conditions. However, this
effect is very unlikely to influence the imagery conditions, for at least two reasons. First,
there were no fixed landmarks in the room to which the subjects could have referred
when setting the indicator to the horizontal. Second, subjects could not easily reproduce from memory the indicator's position because the sequence of tasks within the
perception and imagery conditions was counterbalanced and, prior to the imagery
conditions, there was a break during which the laboratory room was fully illuminated.
Figure 2 summarizes the procedure in all perception and imagery conditions.
Partial
perception

Image
completion

Surround
motion

Complete
perception

Image
generation

Figure 2. Summary of all perception conditions (top row) and imagery conditions (bottom row)
included in this study.

2.3.3 Subjective reports. For each subject, we assessed how difficult he or she found the
imagery tasks by requesting ratings on a five point scale, from 1 (``the mental image
of the moving spots appeared perfectly clear and vivid as during normal vision'') to 5
(``there was no visual mental image at all, I just knew that I was thinking of the
moving spots''). After each condition, we asked the subjects to indicate their experience
of vection on a percentage scale; an estimate of 0 : 100 signified that all the motion
appeared in the display and, in contrast, an estimate of 100 : 0 signified that all the
motion was experienced as body rotation. The subjects were instructed to estimate
the average vection they experienced over the entire duration of the condition. The
percentages given for display rotation (eg 60%) and body rotation (eg 40%) had to add
up to 100. After each condition, the subjects also reported whether they experienced
subjective body tilts when the drum was rotating.

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951

3 Results
3.1 Surround-motion condition
The peripheral visual motion of the randomly positioned spots surrounding the central
portion was present in all perception and imagery conditions. These spots served as a
perceptual crutch when people imagined the pattern moving, and exerted a directional
shift on SVH compared to the SVH measured when the drum was stationary.
Figure 3 shows that all nine subjects who viewed clockwise drum rotation adjusted
the SVH in the clockwise direction, with a mean bias of 7.28 (SD 2.58). The nine
subjects who viewed counterclockwise drum rotation adjusted the SVH in the counterclockwise direction, with a mean bias of 9:28 (SD 2.98). As evident in figure 3, there
was a slight tendency for a stronger bias in the counterclockwise condition (t8 1:828,
p 0:105). Women and men did not differ in their responses to peripheral motion.
A shift in the clockwise direction is expressed with a positive sign and a shift in the
counterclockwise direction is expressed with a negative sign. These influences induced by
surround-motion stimulation were corrected for slight individual SVH deviations from
the physical horizontal when the drum was stationary. These adjustments of the SVH
without drum rotation were very accurate and clustered around a mean of 1.48 (SD 3.78).
This slight deviation was in the clockwise direction from the physical horizontal.
CW
20

Change of SVH=8

15

Rotation CW

10
5
0
5
10
15
Rotation CCW
20
CCW

Figure 3. The mean bias induced by the rotating background on the subjective visual horizontal
(SVH). Black columns represent the bias during counterclockwise rotation whereas white columns
represent the bias during clockwise rotation. Error bars are standard deviations. Slight deviations
from the physical horizontal in stationary conditions are corrected for each subject.

3.2 Modification of the subjective visual horizontal during perception

We computed the perceptual bias by subtracting the mean SVH in the partial-perception
condition (a disk with inner circle consisting of six spots) from the mean SVH in the
complete-perception condition (a disk with both circles consisting of eighteen spots).
The perceptual bias induced by the additional motion cues of the outer band of spots
led to an average increase of 2:18 (SD 2.18) for clockwise drum rotation and 3:58
(SD 5.78) for counterclockwise drum rotation. The presence of the outer band of spots
biased fifteen of eighteen subjects in the direction of drum rotation; that is, the outer
band of spots biased the SVH in a clockwise direction (positive sign) during clockwise
roll stimulation, and biased the SVH in a counterclockwise direction (negative sign)
during counterclockwise roll stimulation. The individual values ranged from 6:18 to
1:48 for clockwise rotation (one out of nine subjects had a negative bias) and
from 14:68 to 3:28 for counterclockwise rotation (two out of nine subjects had a
positive bias).

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F W Mast, A Berthoz, S M Kosslyn

3.3 Modification of the subjective visual horizontal during imagery

For the imagery conditions, the biasing effect was defined as the mean angular
displacement of the indicator compared to the condition in which no spots were visualized. The mean imagery bias was computed by subtracting the mean SVH in the
partial-perception condition from the mean SVH in the image-completion condition
when the twelve spots on the outer band were visualized. In this case, the subjects viewed
exactly the same set of stimuli and the instruction to visualize was the only difference
between the image-completion and partial-perception conditions (see figure 2).
3.3.1 Image-completion condition. The key finding was that visualizing spots in the
imagery conditions deflected the spatial judgments in a very similar manner as
occurred when the spots were actually visible. The bias in imagery correlated strongly
with the bias in perception; the Pearson correlation coefficients for complete perception versus image completion were r7 0:85, p 0:029 for clockwise rotation and
r7 0:73, p 0:002 for counterclockwise rotation.
Figure 4 shows that similar biases were induced when the outer band of spots was
imagined, in contrast to when they were actually viewed, in the complete-perception
condition. The individual biases ranged from 4:48 to 0:28 during clockwise roll
stimulation (one subject had a negative bias) and from 4:08 to 2:88 during counterclockwise roll stimulation (three subjects had a positive bias). All individual biases
in imagery and perception that exceeded 28 were significantly different from partial
perception ( p 4 0:03).
CW
10

Rotation CCW

Rotation CW

Imagery bias=8

6
4
2
0
2
4
6
8
10
CCW 14 1210 8 6 4 2 0 2 4 6
CCW
Perception bias=8

8 10 12 14
CW

Figure 4. The ordinate shows the mean imagery bias on SVH induced by imagining twelve dots
forming the outer circle. The presence of the outer circle exerts a bias on the SVH as shown on
the abscissa representing the perception bias. Both biases are corrected by comparisons with the
rotating pattern with the inner circle, which was visible in imagery and perception.

3.3.2 Image-generation condition. The image-generation condition, in which the subjects

had to visualize all eighteen spots, produced comparable results: the correlation of
the data from image generation versus complete perception was r7 0:90, p 0:0002
for clockwise rotation and r7 0:76, p 0:015 for counterclockwise rotation. In this
case, the perceptual bias was computed by subtracting the mean SVH in the surroundmotion condition (a disk with only the fixation spot on it) from the mean SVH in
the complete-perception condition (a disk with both bands, consisting of eighteen spots).
The mean imagery bias was computed by subtracting the mean SVH in the surroundmotion condition from the mean SVH in image-generation condition when the eighteen
spots on the outer circle were purely imagined (see figure 2).

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953

Furthermore, results from the two imagery conditions generation and completion
were strongly correlated: r7 0:83, p 0:004 for imagined motion in a clockwise
direction and r7 0:75, p 0:017 for imagined motion in a counterclockwise direction. However, the biasing effects were significantly smaller when images of both
bands of spots were generated (t8 2:23, p 0:057 for clockwise imagined motion and
t8 2:65, p 0:029 for counterclockwise imagined motion).
3.4 Subjective reports and latencies of vection
Twelve subjects indicated that the image-generation condition was more difficult than
the image-completion condition. The average rating was 3.7 for generation and 3.1 for
completion (t17 1:91, p 0:0733). The mean vection estimates, which were determined
on a percentage scale, showed an increase of subjective body rotation from 26.9%
(range 0% to 60%) for the surround-motion condition to 33.8% (range 5% to 85%) when
the pattern was imagined in the image-generation condition (t17 2:15, p 0:0462).
The vection induced by image completion, which resulted in 35.7% mean subjective
body motion, did not significantly differ from the partial-perception condition (t17 0:88,
p 0:3917). The majority of subjects, thirteen out of eighteen, reported body tilts
in each of the experimental conditions. Among the subjects who reported subjective
body tilts, seven viewed CW rotation and six viewed CCW rotation. They reported body
tilts to the left when the drum was rotating CW, and, correspondingly, they reported
body tilts to the right when the drum was rotating CCW. The subjects who did
not experience body tilt felt perfectly upright throughout all imagery and perception
conditions.
We found no differences between women and men in any of the imagery or perception
conditions. The mean vection latencies for the five conditions are shown in table 1.
The mental effort to visualize increased the latency for vection onset relative to the
corresponding perceptual condition, in the image-generation condition from 10.4 to
18.9 s t17 2:53, p 0:0214) and in the image-completion condition from 12.8 to 21.2 s
(t17 1:92, p 0:0724). The individual latencies ranged from 2.7 s (in the partialperception condition) to 90 s (in the complete-perception condition).
Table 1. Mean vection latencies and standard deviations (in seconds) for all conditions.
Condition

Latency

Standard deviation

Surround motion (peripheral spots only)

Partial perception (six spots on central disk)
Complete perception (eighteen spots on central disk)
Image completion (six spots on central disk)
Image generation (peripheral spots only)

10.4
12.8
17.3
21.2
18.9

6.3
9.6
19.1
18.6
13.3

4 Discussion
The present results provide evidence that allocentric percepts can be affected by top ^
down processing via visual mental imagery. Moreover, we have demonstrated that
imagined motion can in fact interfere with the spatial perception of allocentric orientation. Previous studies on vection have predominantly focused on investigating the
effects of different displays. For example, Brandt et al (1973) proposed that a display in
the periphery more effectively induces vection than a display in the center of the
visual field, whereas Howard and Heckmann (1989) reported that this superiority
of the periphery may have been confounded with perceived relative distance. To
our knowledge, this debate fails to note potential top ^ down influences, which were
specifically investigated in the current study.

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F W Mast, A Berthoz, S M Kosslyn

The results from both imagery conditions demonstrate that mental images can
`stand in' for perceptual stimuli, and exert the same directional influence on the SVH
as the corresponding perceptual situation. A major requirement in imagery studies
is ensuring that the subjects did in fact use visual mental imagery during the task.
In principle, the effect of visual mental imagery could be mediated by an attentional
effect: in this case, attention would modulate the biasing effect of motion in the
periphery, which was present in all conditions (as illustrated in figure 2). However,
the biases in the two imagery conditions, generation and completion, differed significantly, and thus the observed effect cannot be ascribed simply to increased attention
to motion in the periphery. Moreover, the subjects required more time for vection to
reach its onset in the image-generation condition, which again is not consistent with
the notion that subjects simply allocated their attention to the peripheral surround
motion; if so, the subjects should have been faster in the surround-motion condition
than in the image-generation condition. Lepecq et al (1995) showed that onset latencies can in fact be used to study the role of cognitive effects on vection; they found
prolonged onset latency when subjects were given information that was not consistent
with the effects induced by the vection stimulus.
The fact that the image-generation condition induced a smaller change in SVH
than we found in the image-completion condition may be due to capacity limits in visualizing complex patterns, which could have made it harder to maintain the mental image
of both bands. In addition, the presence of the inner band in the image-completion
condition may have served as `perceptual assistance', facilitating the formation of the
mental image.
Furthermore, the results from this study also showed larger bias when the pattern
rotated counterclockwise, compared to counterclockwise. This effect has been reported
in previous studies by Held et al (1975) and Guerraz et al (1998).
One could also question whether local visual motion contrast effects contributed
to our results. Local visual motion contrast could be perceived when the lasers appear
near one or both of the bands of spots in the perception conditions. Though this could
account for the perceptual bias, imagined motion also exerted a similar bias when
such local contrast cues were absent in the image-generation condition. It could be
argued that the effect in imagery could be mediated by top ^ down activation of a
motion contrast detector operating in a locally confined area of the visual field.
Murakami and Shimojo (1996) have proposed a motion contrast detector with an
antagonistic center ^ surround organization, which is possibly located in area MT.
However, the stimulus configuration in the present study also elicited vection, which
was significantly increased when motion was imagined in the completion task. Therefore,
local motion contrast cannot solely induce the bias produced by the rotating patterns.
In addition, the fact that subjects experienced an increase in vection during visual
mental imagery suggests a possible relation between visual mental imagery and vection
susceptibility. Indeed, individual sensitivity to vection may predict performance in other
cognitive and perceptual tasks. Individuals differ reliably in the strength of experienced
vection (Kennedy et al 1996), and also differ in their abilities to perform various
imagery tasks (Kosslyn et al 1984). However, to our knowledge, even the relation
between vection and specific imagery abilities, such as mental image transformation
abilities, has not been addressed empirically.
Psychophysical studies have provided evidence that the processing of real motion
engages neural mechanisms that are also used when motion is purely imagined, for
example in time-to-contact judgments (Gilden et al 1995) or in mental-rotation tasks
(Corballis and McLaren 1982; Heil et al 1997). And mental-rotation studies have
provided evidence that a stimulus representation is rotated through a trajectory of states,
which correspond to the rotation of an external object (Cooper 1976; Shepard and

Mental imagery and vection

955

Cooper 1982). Moreover, mental rotation has been shown, at least in some conditions,
to engage motor areas of the brain (eg Parsons et al 1995; Kosslyn et al 1998). In the
current study, the subjects were, in fact, continuously rotating a mental representation
of the previously memorized circular arrangement of spots. It is plausible that visualizing a rotary motion pattern in the current task does at least partially share the same
mechanisms that are involved in mental-rotation tasks.
Research in multisensory integration suggests another approach for interpreting
the current results. On the basis of the idea that sensory information can be differentially `weighted' by modality, it is conceivable that the effort to visualize could have
lessened the relative influence of gravity-receptive cues. For example, a neuroimaging
study by Brandt et al (1998) showed that circular vection activates a medial parietooccipital visual area and simultaneously deactivates the parieto-insular vestibular cortex.
The authors suggest a reciprocal inhibitory interaction between visual and vestibular
information, which could reduce perceptual ambiguities. In the current context, our
findings could be interpreted as an extension of Brandt et al's (1998) view; imagined
motion could activate the same mechanisms that are also used in the processing of
visual motion, and thereby inhibit vestibular information. Alternatively, Mittelstaedt's
(1991) theory suggests another interpretation for our imagery effects. His approach
relies on a body-centered reference, the idiotropic vector, which is independent of
gravity and points into the direction of the body axis. In the current context, this
theory would predict an increased visual influence of the rotating field as a consequence of a diminished idiotropic influence. To our knowledge, no study has been
conducted that actually shows that top ^ down processing, such as visual mental
imagery, can attenuate the idiotropic gain. Yardley's (1990) clinical findings with
proprioceptive deficits have demonstrated that changes in idiotropic gain could in fact
play a role in allocentric orientation tasks.
In summary, our results demonstrate that imagined motion can exert the same
directional influence on allocentric orientation judgments as the perception of the
corresponding stimulus. The results clearly show that the mechanisms underlying the
processing of motion information are susceptible to top ^ down influences. Knowledge
about how top ^ down processes are used in spatial tasks offers a potential bridge
between higher cognitive processes and cortical mechanisms involved in sensory integration.
Acknowledgements. This work was supported by NIH grant R01 MH60734-01. Fred Mast is
supported by a grant from the Swiss National Science Foundation. We are grateful to the Man
Vehicle Laboratory at MIT for providing technical support.
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