Binocular contributions to linear vertical vection

Robert S. Allison
#
$
Department of Electrical Engineering and Computer
Science,
York University, Toronto, Canada
April Ash $
Department of Electrical Engineering and Computer
Science,
York University, Toronto, Canada
School of Psychology, University of Wollongong,
Wollongong, Australia
Stephen Palmisano
#
$
School of Psychology, University of Wollongong,
Wollongong, Australia
Compelling illusions of self-motion, known as vection,
can be produced in a stationary observer by visual
stimulation alone. The role of binocular vision and
stereopsis in these illusions was explored in a series of
three experiments. Previous research had provided
evidence of stereoscopic enhancements for linear
vection in depth (e.g., Palmisano, 1996, 2002). Here we
examined for the first time the effects of binocular vision
and stereopsis on linear vertical vection. Vertical vection
was induced by the upward or downward translation of
large stereoscopic surfaces. These surfaces were
horizontally oriented depth corrugations produced by
disparity modulation of patterns of persistent or short
lifetime dot elements. We found that binocular viewing
of such surfaces significantly increased the magnitudes
and decreased the onset delays of vertical vection.
Experiments utilizing short lifetime dot stereograms
demonstrated that these particular binocular
enhancements of vection were due to the motion of
stereoscopically defined features.
Introduction
Self-motion through the world generates patterns of
visual stimulation called optic ow (Gibson, 1950).
Most self-motion research to date has considered the
optic ow provided to a single eye; however, self-
motion (like object motion) produces different patterns
of optic ow at the left and right eyes. This raises the
following question: Is there an additional, or even
independent, contribution of binocular viewing or
binocular motion stimulation to the perception of self-
motion? In this paper we consider this question for
lamellar ow, corresponding to the simulated vertical
motion of either the self or the whole visual environ-
ment (i.e., self- or scene-motion parallel to the coronal
plane of the head). In addition to uncovering the
underlying sensory processing of self-motion, the
answer to this general question is practically important
for self-motion simulation applications such as vehicle
simulators, rides, and virtual reality, where the benet
of stereoscopic display must be weighed against the
cost and technical challenges of presenting full-eld
stereoscopic displays.
There are several ways that binocular vision could
contribute to the perception of linear self-motion, over
and above the view provided by a single eye: (a)
increased eld of view; (b) binocular summation; (c)
stereoscopically dened features and motion; (d)
improved information about environmental layout,
depth, and parallax; and (e) improved perception of
rigidity and structure. Here, for the rst time, we
describe each of these possible binocular contributions
in detail and then discuss the available evidence
supporting their involvement in the perception of self-
motion.
Field of view
One of the most basic benets of binocular vision is
the increased eld of view provided, an advantage
maximized in lateral-eyed animals but important even
Citation: Allison, R. S., Ash, A., & Palmisano, S. (2014). Binocular contributions to linear vertical vection. Journal of Vision,
14(12):5, 123, [Link] doi:10.1167/14.12.5.
Journal of Vision (2014) 14(12):5, 123 1 [Link]
doi: 10. 1167/ 14. 12. 5 ISSN 1534-7362 2014 ARVO Received March 17, 2014; published October 9, 2014
for frontal-eyed animals, such as humans, that typically
have a signicantly larger total eld of view than can be
seen from either eye. Both circular and linear vection
reportedly increase with stimulus area (e.g., Telford &
Frost, 1993), so the increased binocular eld of view
should promote vection. We do not consider this aspect
further as stimuli in the present study were restricted to
only the binocular visual eld and were visible to both
eyes.
Binocular summation
Binocular summation refers to the combination of
typically redundant signals from the left and right eyes
to form a stronger signal, improve task performance, or
improve the signal-to-noise ratio (for a recent review
see Howard & Rogers, 2012). Increases in effective
contrast from binocular summation improve the
detectability and discriminability of low-contrast stim-
uli but not high-contrast stimuli (Legge, 1984) and
performance on spatial vision tasks at low contrast
(Banton & Levi, 1991; Home, 1978). Thus, binocular
summation could enhance the visibility of low-contrast
stimuli and allow them to contribute to vection. In line
with this reasoning, binocular summation has been
shown to improve global motion coherence detection at
low contrast (Hess, Hutchinson, Ledgeway, & Man-
souri, 2007).
Binocular summation of similar inputs is not the
only way that binocular inputs can combine. The most
important form of combination of dissimilar inputs is
stereopsis, which is considered below; however, differ-
ent information to the two eyes can also be combined
to form a composite dichoptic image that is the sum of
the monocular images. Such binocular combination of
dissimilar images is most likely at low contrast;
binocular rivalry is more likely at higher contrast (Liu,
1992). Hess et al. (2007) also found that dichoptic
presentation (signal dots to one eye and noise dots to
the other) did not improve performance over monoc-
ular presentation (signal and noise dots both presented
to one eye; a uniform gray eld of mean luminance
presented to the other). When combined with their
ndings on contrast dependence, these authors con-
cluded that (a) the motion signals for the two eyes are
combined at an early, contrast-dependent stage of
processing and (b) global motion processing is purely
binocular.
Stereoscopically defined features and motion
Stereopsis is a potent cue to depth and consequently
moving stereoscopic features are a cue to three-
dimensional motion. Since the two eyes are laterally
separated in the head, the images on the two retinas
differ. In particular, the position of the image of an
object can differ on the two retinas; this is referred to as
a positional disparity. In the 19th century, Wheatstone
(1838) demonstrated clearly that a compelling sense of
depth perception, known as binocular stereopsis, could
be obtained from these binocular disparities. When an
observer moves in depth relative to an object its
positional disparity changes, providing the changing
disparity cue to motion-in-depth; furthermore, when an
observer or object moves in depth the images of the
object typically move at different velocities in the two
eyes, leading to the cue of interocular velocity
difference (Regan, 1993). Similarly, disparity change
and interocular velocity differences can provide infor-
mation about the direction of motion in depth (Regan,
1993). Psychophysical studies have shown that motion
in depth can be elicited by either changing disparity or
interocular velocity differences (Allison & Howard,
2011; Allison, Howard, & Howard, 1998; Shioiri,
Saisho, & Yaguchi, 2000). During self-motion these
interocular velocity difference and changing disparity
cues are available across the optic ow eld. These
binocular cues could elicit or contribute to perceived
motion in depth and hence to linear vection in depth.
Palmisano (2002) found that adding consistent stereo-
scopic depth to simulated forward self-motion displays
increased apparent self-motion speed and distance
traveled. Additional experiments suggested that the
enhanced vection was due to binocular cues to motion
in depth rather than improved perception of layout of
the three-dimensional scene.
Binocular disparity can also dene stereoscopically
dened contours that are not visible in the monocular
images. Such features are known as cyclopean features
because they are dened and visible only after the
binocular inputs are combined (Julesz, 1971). Cyclo-
pean images can support the perception of form and
motion and even geometrical illusions. If the position
of the cyclopean contours is changed over time then the
cyclopean form appears to move, and this stereoscop-
ically dened motion could produce vection. To our
knowledge the possibility that moving cyclopean
contours can induce vection has never been tested.
However, moving cyclopean contours in a dynamic
random-dot stereogram can produce optokinetic nys-
tagmus in observers with normal stereoscopic vision
(Fox, Lehmkuhle, & Leguire, 1978). Furthermore,
Loomis and colleagues have reported that observers
can make accurate judgments about the direction of
self-motion (i.e., heading judgments; Macuga, Loomis,
Beall, & Kelly, 2006) and guide interceptive movements
(Loomis, Beall, Macuga, Kelly, & Smith, 2006), even if
the only information about the simulated self-motion is
provided by the motion of cyclopean features.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 2
Information about environmental layout, depth,
and parallax
The motion perspective in optic ow is complex and
potentially ambiguous. One problem is the scale
ambiguity of the motion perspective in the ow eld
did the ow result from a large motion in a large
environment or a small motion in a small environment?
Binocular information about distance from vergence
and vertical disparity could, in principle, provide the
scaling factor, at least in relatively near space (Rogers
& Bradshaw, 1995). Butler, Campos, B ulthoff, and
Smith (2011) found that, for stationary observers,
heading discrimination was not improved when ste-
reoscopic cues were added to optic ow displays
simulating self-motion (compared with when both eyes
viewed an equivalent nonstereoscopic display). How-
ever, combined visualvestibular just-noticeable differ-
ences for physically moving observers were closer to
optimal cue integration under stereoscopic presenta-
tion; suboptimal combined just-noticeable differences
were obtained under binocular, nonstereoscopic pre-
sentation. As the task was simply discrimination of the
heading direction relative to straight ahead, the
assumed scale of the optic ow display should have
affected only the apparent magnitude, not the direc-
tion, of the heading offset. The authors speculated that
disparity cues help determine the ambiguous scale of
the optic ow display and thus promote integration
with the vestibular cuesthis assumes that effective
visualvestibular integration depends on compatible
vestibular and visual estimates of heading direction.
Another potential ambiguity arises from difculties
in parsing the ow eld into various components. Local
ow can arise from self-motion or object/scene motion.
Even if object/scene motion can be discounted, the
effects of linear and rotational self-motion also need to
be distinguished. Theoretically (Longuet-Higgins &
Prazdny, 1980) monocularly available optic ow could
be so decomposed, assuming both rigidity and depth
variation in the scene. However, this processing would
be simplied substantially if the layout and depth
relations in the environment could be estimated.
Consistent with this notion, van den Berg and Brenner
(1994) reported that heading judgments were more
tolerant to randomly directed local motion noise when
short-duration (1.5 s) looming optic ow was presented
stereoscopically as opposed to synoptically (i.e., iden-
tical images were presented to the two eyes). Interest-
ingly, tolerance to this motion noise was similar when
the moving dots had a xed disparity (appropriate to
their simulated three-dimensional layout on the rst
frame) compared with when their changing disparities
represented the appropriate motion in depth. This
latter nding suggested that the improved resilience was
due to information about spatial layout rather than
stereoscopic cues to motion in depth. As noted in the
previous paragraph, Butler et al. (2011) did not nd
any stereoscopic advantage for heading perceptions
based on purely visual self-motion simulation.
The only systematic studies of the effect of stereo-
scopic layout cues on vection were conducted by
Palmisano (1996, 2002). In the earlier paper, he found
that stereoscopic displays elicited stronger (shorter
latency and longer duration) vection in depth than did
monocular or nonstereoscopic binocular displays. He
considered that the stereoscopic enhancement could
have been due to improved information about the
layout of the simulated objects, improved impressions
of depth, or improved motion-in-depth cues. Since the
monocular displays contained strong cues to relative
depth, he argued that it was unlikely that stereoscopic
cues could further help disambiguate layout. In the later
paper, he found that stereoscopic information did not
appear to be improving vection in depth by increasing
the perceived maximum extent of displays or by making
displays appear more three-dimensional. This led him to
conclude that the stereoscopic enhancement of vection
was due to stereoscopic motion-in-depth cues (rather
than an improved perception of layout). In the present
experiments we simulated vertical motion, rather than
motion in depth, in order to focus on aspects of the
binocular contribution to vection other than binocular
motion in depth.
Improved perception of rigidity and structure
Grigo and Lappe (1998) showed that transparently
superimposing a horizontally translating optic ow
eld on a looming ow pattern shifts the perceived
focus of expansion (FOE). Lappe, Bremmer, and van
den Berg (1999) suggested that this shift results from
compensatory mechanisms that interpret the hori-
zontal translation as a headeye rotation and correct
the observers heading estimate accordingly. Accord-
ing to their proposal, visual motions of the most
distant parts of the scene provide the best estimates of
head-and-eye rotation (as the looming and parallax
arising from whole-body self-motion is more evident
in the image motions of nearer points). Importantly,
they also found that the shift in the apparent FOE was
modulated by the relative disparity between the
translating and looming patterns. The weakest shift in
the FOE was found when the translating pattern was
stereoscopically simulated to be in front of the
looming patternpossibly because this particular
arrangement of motion vectors implies a nonrigid
optic ow eld, which would be expected to degrade
the perception of self-motion.
Depth information has been identied as being
important for the effective visual parsing of object
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 3
motion from self-motion (Warren & Rushton, 2009).
Rigid movement of all objects in the world is rare and
typically results from self-motion rather than the
movement of the world. There is evidence that visual
environments that are perceived as rigid are more likely
to induce vection or postural responses than are those
not seen as rigid (Holten, Donker, Verstraten, & van
der Smagt, 2013; Nakamura, 2010; but see Palmisano,
Allison, & Howard, 2006; Palmisano, Kim, & Free-
man, 2012). However, the role that binocular disparity
plays in perceived rigidity has not been explored.
Consistent disparity and optic ow-based information
about layout should promote the perception of self-
motion through a stable, rigid environment. By
contrast, stereomotion or stereoscopic depth inconsis-
tent with the optic ow should further reinforce the
perception of object movement or deformation.
Related but distinct from rigidity, the foreground
background relationship of objects moving in the image
is important. Binocular disparity is potentially impor-
tant to separate foreground from background and to
provide depth order. Logically, a moving background
should signal self-motion, whereas foreground objects
could move due to either object motion or self-motion.
The more distant surface is usually perceived as
background, and vection is usually perceived in the
direction opposite to the background motion (Ohmi,
Howard, & Landolt, 1987). Similarly, Ito and Shibata
(2005) superimposed either (1) radially contracting and
expanding optic ow or (2) two separate expanding
optic ows that were horizontally shifted (i.e., the two
FOE were offset). They found that when separated in
depth (based on disparity), vection direction was
always based on the more distant ow. Nakamura
(2008) used binocular disparity to promote different
foregroundbackground relations between central and
peripheral stimuli. He found that central and peripheral
stimuli covering equal retinal area produced equivalent
vection when the foregroundbackground relationship
was controlled. He concluded that previous evidence
for peripheral dominance in vection was due to
confounding of apparent depth and eccentricity.
The current study: Stereoscopic contributions to
linear vection
While there are many potential benets that binoc-
ular vision could provide to self-motion perception, the
role of binocular vision in vection has received very
little empirical examination. What little research has
been conducted has focused on vection in depth, and
the binocular advantage found for this has been
suggested to arise from the extra stereoscopic infor-
mation about motion in depth. Whether binocular
vision also improves other types of self-motion is
currently unknown. Furthermore, there are many other
ways that binocular vision could improve self-motion
perception. Here we investigate (a) whether binocular
vision can improve a very different type of vection (not
involving self-motion in depth but rather self-motion
parallel to the frontal plane) and (b) whether binocular
benets (other than added information about motion in
depth) might also signicantly enhance vection.
Due to the close link between optic ow and the
environmental layout during self-motion, we expected
that linear vection would vary depending on the cues to
three-dimensional layout present in the scene. In this
study we concentrated on the contribution of binocular
vision and stereopsis to the perception of vertical linear
vection induced by lamellar ow. Simulating transla-
tional self-motion parallel to a frontal cyclopean
surface permits assessment of the role of binocular
viewing, stereoscopic surface perception, and stereo-
scopic features in the production of vection independent
of binocular cues to self-motion in depth (the so-called
stereomotion cues).
The current experiments examined the ability of
cyclopean motion stimuli (vertically translating, dis-
parity-dened depth corrugations) to induce vection.
We examined not only whether cyclopean motion
stimulation could enhance the vection induced by
monocular motion signals but also whether such
stimulation induces vection on its own. We also
examined the vection induced by binocular viewing
both with and without stereoscopically-dened depth
corrugations and manipulated the strength of the
monocularly available motion signals. To investigate
these questions systematically we (a) compared the
vection induced by dynamic random-dot stereogram
(DRDS) with random-dot stereogram (RDS) stimuli
to isolate the effects of cyclopean motion (the former
providing only cyclopean motion information; the
latter providing both cyclopean and monocular
motion information); (b) varied the three-dimensional
stimulus shape (i.e., the disparity waveform) to
control the presence and extent of stereoscopically-
dened moving features; (c) evaluated the inuence of
stereoscopic vision by comparing vection elicited in
binocular viewing with monocular viewing of the same
stimuli; (d) evaluated the effect of moving cyclopean
form by comparing the vection induced by moving
disparate surfaces (i.e., with depth corrugations) with
that induced by zero-disparity (i.e., at) surfaces; and
(e) controlled the strength of monocular motion
signals in our vection-inducing displays by varying the
lifetime of moving features. We also varied the
simulated speed of self-motion and compared free
viewing with xation to test the generality of our
results across both stimulus factors (known to be
important in the production of vection; for review see
Howard, 1982).
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 4
General methods
The experiments were performed under a protocol
adhering to the Declaration of Helsinki and approved
by the Human Participants Review Committee of York
University.
Stimuli were presented on a large stereoscopic
television monitor (LG 55L W5700, LG Electronics,
Seoul, Korea). This display has a lm-patterned
retarder overlay that acts as a micropolarizer array to
circularly polarize the light. The pitch of the micro-
polarizer array matches the display pitch and is
overlaid on the liquid-crystal display pixel grid so that
the circular polarization of light from even rows of
pixels is opposite the circular polarization of light from
odd rows. Stereoscopic presentation was achieved by
presenting the right-eye image on odd rows and the left-
eye image on even rows. The observer wore glasses with
an orthogonal pair of polarizing lters that matched
the micropolarizer array. These glasses act as an
analyzer and ensured that the left eye saw only the even
rows and the right eye saw only the odd rows. The level
of interocular crosstalk (white on black) was 0.4%. The
images were presented at the native resolution of the
display (1920 1080 pixels) and refreshed at 60 Hz.
The subject was seated in a chair and viewed the
screen at a distance of 215 cm, with the chair adjusted
to center the point midway between the eyes with the
screen. Each pixel subtended 1.0 arcmin. At this
distance the screen subtended 31.48 wide by 18.08 high
(121 68 cm), but a rectangular aperture was placed
141 cm from the screen and dened the stimulus size at
27.38 by 17.68. Two vertical bars subtending 0.778 in
width and extending across the aperture were located
10.88 to the left and right of middle of the aperture.
This provided the strong impression of looking through
a window at the display beyond (similar to looking out
the window of a moving vehicle). This manipulation
generated very compelling vection illusions despite the
relatively small central display. Note that compelling,
centrally induced vection has previously been reported
for both lamellar and radial optic ow when these
stimuli are viewed through such apertures (e.g.,
Andersen & Braunstein, 1985; Andersen & Dyre, 1989;
Howard & Howard, 1994; Ohmi & Howard, 1988;
Telford & Frost, 1993). The room was light proofed,
and an enclosure covered in black cloth and cardboard
ensured that only the display was visible within the
dark frame of the aperture window.
The stimuli were computer generated on a desktop
workstation (Intel i7-860 2.80 GHz 4 GB, Intel Corp.,
Santa Clara, CA; Windows 7 64 bit, Microsoft Corp.,
Redmond, WA; Nvidia GE-Force GTX460, Nvidia
Corp., Santa Clara, CA). Stereoscopic image sequences
were produced and presented with Python scripts using
OpenGL (via pyglet 1.1.4; [Link]). Participant
responses were recorded with a Logitech dual-action
gamepad (Logitech International SA, Morges, Swit-
zerland).
Stereoscopic image sequences each depicted a hori-
zontally oriented, depth-modulated waveform (visible
only during binocular viewing). The disparity prole of
each waveform varied as a function of vertical screen
position according to frequency (f) of 6 cycles per screen
and peak disparity (D
peak
) of 5.44 arcmin (10.88 arcmin
peak to peak). The disparity prole was consistent with
a sinusoidal wave, a sawtooth wave, a triangle wave, or a
square wave surface in depth, in each case centered on
the screen disparity (Figure 1). When the stimulus
moved, the waveform translated up or down on the
screen (depending on the sign of the velocity).
Each frame of the motion stimulus comprised 4,000
antialiased dot pairs (dot diameter 2.0 arcmin; dot
luminance 73.9 cdm
2
on a 0.007 cdm
2
background),
randomly positioned in the stimulus image but with a
matching or correlated position in images of both eyes.
The disparity required for a given dot pair was
introduced by shifting the dots in equal and opposite
directions in the two eyes by half of the required
disparity. Three types of RDS motion stimuli (which
varied in terms of dot lifetime and the types of motion
that were available) were examined:
(1) RDS motion stimuli. These displays consisted of
left- and right-eye dot pairs, which were created on
the rst frame and persisted until the end of the
stimuli. Each dot pair had an initial two-dimen-
sional image position and disparity for the rst
frame. As the waveform translated on subsequent
frames, the disparity and horizontal position of the
dot pair remained the same but its vertical position
was updated so that all dots moved upward or
downward together with the disparity wave (as
though the dots were painted on the moving three-
dimensional surface).
(2) DRDS motion stimuli. In these displays the
positions of all left- and right-eye dot pairs were
refreshed every 16.67 ms. Essentially, 4,000 new dot
pair positions were chosen on each frame, and the
disparity of these dot pairs was assigned according
to the current position of the disparity waveform.
(3) Limited-lifetime RDS stimuli. These displays were
similar to DRDS except the left- and right-eye dot
pairs persisted over several frames, after which they
were extinguished and their positions were refreshed
(as in the DRDS). Essentially, 4,000/L new dot pair
positions were chosen on each frame for the
expiring dots, where L is the dot lifetime in frames,
and the disparity of these dot pairs was assigned
according to the current position of the disparity
waveform.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 5
Figure 1. (a) Disparity waveforms used in the experiment (vertically offset for visibility). Disparity was modulated as a function of
vertical (y-axis) position on the screen. Disparity modulations had the same peak amplitudes and period but differed in the smooth
and discontinuous disparity changes. (b) The subject viewed the moving corrugated surface through an aperture in a dark room.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 6
All of these motion stimuli were presented for 30 s
(1,800 frames). Representations of RDS and DRDS
stimuli are shown in Movie 1 and Movie 2. For trials
where xation was controlled, a xation cross was
presented in the center of the display at zero disparity
with respect to the screen. The cross consisted of one
horizontal and one vertical line, each 21.8 arcmin long
with a stroke (width) of 5.4 arcmin, visible in both eyes.
The observers task was to monitor their sensation of
self-motion and rate its strengththat is, how com-
pelling the experience wasrelative to a standard using
magnitude estimation (Stevens, 1975). The standard
stimulus was a reliable vection-inducing stimulus of
moderate strength. The standard was chosen to be
similar to the stimuli presented in the experiment
(differed by experiment; see below) and was presented
to the observer at the beginning of each block of trials.
Subjects were told to (a) assign this standard stimulus a
strength of 50 (the modulus) and (b) make estimates
of vection strength proportionally relative to this
modulus. For instance, if the subjects experience of
vection was twice as strong as the standard they were to
report 100, whereas if vection was only half the strength
of the standard they were to report 25.
In addition to making poststimulus magnitude re-
sponses, subjects were instructed to continuously mon-
itor whether they experienced vection during each trial. If
they experienced vection at any time they were told to
press a gamepad button and hold it while the sensation
was maintained, releasing the button whenever the sense
of vection was lost. From these data we measured the
total duration of the vection sensation during the trial
and the latency of vection. The latency was dened as the
time between the start of the trial and the rst press, or
the trial length (30 s) if the button was not pressed.
Experiment 1: Can vection be
induced or enhanced by cyclopean
motion signals?
The primary purpose of Experiment 1 was to
determine whether vection could be induced by purely
cyclopean display motion. To this end we studied the
perception of vection elicited by two types of binocularly
viewed stereoscopic motion stimuli: one dened only by
Movie 1. Representation of an RDS moving surface. This movie
represents the class of stimulus used but is not a representation
of the actual stimulus. For better tolerance to video
compression and as a web demo, the stimulus has been
modified to a smaller window size, different aspect ratio, fewer
dots, larger dots, and anaglyph presentation. Note that vection
is not likely to be elicited when viewed on a monitor with other
stationary features in view. (Movie 1 can be viewed in the
Supplemental data.)
Movie 2. Representation of a DRDS moving surface. This movie
represents the class of stimulus used but is not a representation
of the actual stimulus. For better tolerance to video
compression and as a web demo, the stimulus has been
modified to a smaller window size, different aspect ratio, fewer
dots, larger dots, and anaglyph presentation. Note that vection
is not likely to be elicited when viewed on a monitor with other
stationary features in view. (Movie 2 can be viewed in the
Supplemental data.)
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 7
cyclopean features and the other dened by both
cyclopean features and monocularly visible texture
motion. Moving RDS and DRDS stimuli provide
different types of information about motion in the
stimulus. With RDS stimuli, the disparity-dened stimuli
move but so do the monocularly visible texture elements
forming the pattern. Thus, RDS stimuli provided both
cyclopean and monocular motion signals indicating
vertical self-motion. In a DRDS, the only coherently
moving features correspond to cyclopean features
dened by change in depth (there were binocular, but no
monocular, motion signals indicating vertical self-mo-
tion; dot lifetime was one frame only). Because the depth
modulation in a DRDS carries all the motion informa-
tion, the form of the modulation might be important.
Square wave modulations provide cyclopean motion
information only at the transitions, whereas continu-
ously varying modulations, such as a sinusoidal wave-
form, provide this information along the entire stimulus
waveform. In this experiment we addressed the ability of
cyclopean stimuli to induce vection.
Methods
See General methods for a description of the
apparatus, stimuli, and procedures.
A total of 12 nave observers (ve males and seven
females ranging in age from 22 to 49 years; mean age 28
6 7.8 years) participated in Experiment 1.
Viewing was always binocular in this experiment.
Trials were grouped into blocks of eight, and the
standard stimulus was presented before each block.
Subjects participated in two sessions, each consisting of
64 trials (plus standards); the trials were counterbal-
anced across sessions and randomized within blocks for
each subject. The standard stimulus was a sawtooth
RDS disparity wave moving at 0.082 m/s with xation
on a central xation cross. As described in the General
methods, observers were instructed to assign a magni-
tude of 50 to this stimulus. Presentation of the standard
stimulus was followed by the experimental trials. The
independent variables were (a) the type of disparity-
dened waveform (square, sinusoid, triangle, saw-
tooth), (b) stimulus speed (0.082 or 0.163 m/s), (c)
disparity direction (i.e., a phase of 08 or 1808 for the
modulating waveform corresponding to whether a peak
or valley was rst seen in the center of the screen), (d)
xation (xation or free viewing), and (e) stereogram
type (RDS or DRDS).
Results
Vection strength rating data
Mean vection magnitude estimates in Experiment 1
are shown in Figure 2. It can be seen that RDS stimuli
(lled symbols) elicited much more robust vection
responses than did DRDS stimuli (open symbols),
which elicited little or no vection. There was very little
Figure 2. Vection magnitude ratings in Experiment 1 as a function of disparity modulation waveform, dot lifetime (stereogram type:
RDS or DRDS), and stimulus velocity. Each data point represents the mean of 12 subjects for the given condition; error bars indicate
61 standard error of the mean.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 8
difference between the vection produced by the different
stimulus patterns. Vection increased with stimulus speed
in the RDS conditions but not in the DRDS conditions.
In the latter case, vection responses were weak or absent
in all conditions, which may explain the lack of any
modulation of the effect by speed.
A repeated-measures analysis of variance (ANOVA)
was used to analyze the results. The dependent measure
was the vection magnitude (strength ratings) and the
independent variables were the factorial combination
of waveform, speed, and lifetime plus disparity
direction and xation type. Tests of univariate hy-
potheses were corrected with Greenhouse-Geisser
adjustments where appropriate.
Stereogram type had the largest inuence on vection
magnitudes, which were considerably larger on average
for RDS stimuli than for DRDS stimuli, F(1, 9)
125.26, p , 0.001, g
2
p
0.933; mean difference M
57.201, 95% condence interval (CI) [45.64, 68.76].
Across subjects and the other factors, vection was
reported on more than 99% of the RDS (unlimited dot
lifetime) trials, while subjects did not experience vection
on 47% of the DRDS (one-frame dot lifetime) trials.
This difference was conrmed by logistic regression
(Wald v
2
52.27, p , 0.001). When vection was
induced, it tended to be much weaker for the DRDS
stimuli compared with the RDS stimuli. Thus, robust
vection was reliably obtained under RDS, but not
under DRDS, conditions.
There was also a signicant effect of stimulus speed,
F(1, 9) 35.93, p , 0.001, g
2
p
0.800, but this main
effect was marginal to a stimulus speed by stereogram
type interaction, F(1, 9) 19.29, p 0.002, g
2
p
0.689.
Vection was larger for the 0.163 m/s stimuli compared
with the slower 0.082 m/s stimuli under RDS stimuli,
mean difference M15.72, 95% CI [9.23, 22.21]. Little
vection was reported under either speed condition for
the DRDS stimuli.
There were no other signicant interactions and no
signicant effects of disparity direction, F(1, 9) 0.602,
p 0.809, g
2
p
0.007; xation, F(1, 9) 1.597, p
0.238, g
2
p
0.151; or waveform type, F(2.07, 18.603)
0.953, p 0.406, g
2
p
0.096 on the vection magnitudes.
Vection time course data
We also measured vection onset times (dened as the
latency from the start of the motion stimulus until the
button press indicating that the subject was experienc-
ing vection) and vection durations (dened as the
accumulated time that the subject pressed the button
during the trial; see Figure 3).
Consistent with the vection strength ratings, RDS
stimuli (lled symbols) elicited vection that persisted
for most of the 30-s trial (median durations of greater
than 25 s), while DRDS stimuli (open symbols) elicited
vection with median durations of less than a few
seconds. There was very little difference between the
vection produced by the different stimulus patterns
Figure 3. Vection duration in Experiment 1 (N12) as a function of stereogram type (RDS or DRDS), fixation condition, and stimulus
velocity. Each data point corresponds to the median across observations; error bars indicate 95% confidence intervals for the
medians.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 9
(disparity modulation type). Vection duration appeared
to increase slightly with stimulus speed and with
xation compared with free viewing.
The onset latency data were subject to pronounced
ceiling effects (onset could not be larger than trial
length), especially for the DRDS conditions where
median onset latency was at least as long as the trial.
Therefore, we analyzed the data using repeated-mea-
sures censored regression (using package censReg in R;
[Link] of onset la-
tency on xation, speed, and lifetime. There was a
signicant effect of stereogram type, t(9) 29.14, p ,
0.001, d 19.43: Mean vection onsets were signicantly
longer for DRDS stimuli (17.96 6 2.95 s, mean 6
standard error of the mean [SEM]) than for RDS stimuli
(4.33 6 0.88 s). Onset was slightly shorter with xation
compared with free viewing, mean difference 1.1 s, t(9)
2.73, p0.023, d1.82, and only slightly longer, t(9)
2.19, p 0.057, d 1.46, for the 0.082 m/s condition
(11.61 6 1.63 s) compared with the 0.163 m/s condition
(10.68 6 1.74 s). There were no other signicant main
effects or interaction effects on vection onset.
The longer the onset latency, the shorter the possible
vection duration for a xed-length trial. Thus, as might
be expected, there was a strong negative correlation
between vection onset and vection duration, r 0.84,
95% CI [0.856, 0.826]. The lack of perfect correla-
tion is due to vection dropouts, where vection ceases
after it has been initiated. Censored regression analysis
demonstrated signicant effects of stereogram type, t(9)
43.84, p , 0.001, d 29.2; xation, t(9) 3.66, p ,
0.01, d 2.44; and stimulus speed, t(9) 3.22, p 0.01,
d 2.15. Consistent and complementary to the vection
onset data, vection duration was increased by higher
stimulus speeds and sustained dots (i.e., RDS as
opposed to DRDS) and was slightly longer with
subjects xating compared with free viewing (Figure 3).
Discussion
The above ndings suggest that vection processing is
relatively insensitive to purely cyclopean motion
stimuli. The DRDS stimuli did not reliably produce
vection responses, mean vection magnitudes were
small, and vection took longer to develop for DRDS
stimuli than for RDS stimuli. This result was found
despite the fact that the DRDS stimuli produced strong
impressions of both a surface modulated in depth and
grating motion. The lack of any disparity waveform-
type effect was not surprising given that we had
expected this inuence to be more apparent for DRDS
(as opposed to RDS) stimuli, and, as it turned out, this
type of stimuli did not elicit much vection.
To maximize the likelihood of obtaining vection we
chose a DRDS stimulus that produced strong depth
and motion impressions. The period of the disparity
modulation was near the peak of the cyclopean depth
modulation sensitivity function (Schumer & Julesz,
1984; Tyler, 1975). Similarly, we used a depth
modulation large enough to provide a strong impres-
sion of depth but modest enough to remain comfort-
ably within Panums fusional limit (Ogle, 1950). While
we varied disparity waveform amplitude, direction, and
frequency over a range of values in pilot experiments,
we did not observe evidence of large sensitivity to these
factors; this is consistent with Kohly and Regans
(1999) nding that observers can ignore variations in
spatial frequency, temporal frequency, and displace-
ment when judging the speed of moving cyclopean
gratings. However, we did not explore this large
stimulus space extensively and thus it is possible that
another cyclopean stimulus could be more effective for
the production of linear vection.
To further promote vection we used a stationary
foreground reference to provide both relative motion
and a frame of reference for vection. (This physical
enclosure and aperture also served the function of
blocking the view of the observers physically station-
ary surroundings.) Like luminance-dened motion,
disparity-dened relative motion is easier to detect and
discriminate than is absolute motion, although there
has been little systematic study of this issue.
In the case of the DRDS stimuli, all of the moving
features were cyclopean. As noted above they were
dened only where the disparity changed. The failure to
nd compelling vection with DRDS stimuli in this
experiment does not necessarily mean that cyclopean
features cannot contribute to vection when combined
with monocularly visible motion signals (as was the
case with the RDS stimuli). In the next experiment we
investigated whether the addition of moving cyclopean
features and binocular input provide additional stim-
ulus for linear vection when combined with monocular
motion signals.
Experiment 2: Binocular versus
monocular viewing of RDS displays
All of the vection-inducing conditions tested in the
rst experiment were binocularly viewed random-dot
stereogram motion displays (both DRDS and RDS).
Only RDS motion displays were found to induce
compelling vertical vection. Here, in Experiment 2, we
compared binocular and monocular viewing of RDS
motion to see whether the binocular viewing of such
displays actually enhances vertical vection. These RDS
motion displays simultaneously present moving cyclo-
pean forms and moving texture under binocular viewing
conditions but only moving textured images under
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 10
monocular viewing conditions. If stereoscopically de-
ned features contribute to visually induced experiences
of self-motion, then we expect that vection will be elicited
more strongly from binocularly viewed RDS motion
stimuli than from monocularly viewed RDS motion.
Methods
See General methods for a description of the
apparatus, stimuli, and procedures.
A total of 15 nave observers (ve males and 10
females ranging in age from 19 to 46 years; mean age 27
6 7.0 years) participated in Experiment 2.
Trials were grouped into blocks of eight trials, with
the standard stimulus presented before each block. The
standard stimulus was a sawtooth RDS disparity wave
moving at 0.082 m/s with xation on a central xation
cross. As described in the General methods, observers
were instructed to assign a magnitude of 50 to this
stimulus. The independent variables were (1) the type
of disparity waveform (square, sinusoid, triangle, or
sawtooth, which was perceptible only when the
stimulus was viewed binocularly), (2) stimulus speed
(0.082 or 0.163 m/s), (3) xation (central xation or free
viewing), and (4) viewing type (binocular or monocu-
lar), with two repeats of every combination for each
observer. The inducing stimuli were always RDS
images moving upward. Monocular viewing (when
required) was achieved by placing an eye patch over the
subjects left eye. There were two experimental sessions,
each session consisting of two blocks under binocular
viewing and two blocks under monocular viewing,
producing a total of 64 trials per subject. The order of
monocular and binocular blocks was randomized and
counterbalanced across sessions for each subject.
Results
Vection strength rating data
Mean vection magnitude as a function of viewing
condition (binocular or monocular), disparity modu-
lation type, and stimulus speed is shown in Figure 4.
The manipulation of most interest in this experiment
was viewing condition. Figure 4 shows that binocular
viewing of disparate moving stimuli produced stronger
mean vection ratings than monocular viewing for all
conditions. Consistent with the results of Experiment 1,
vection magnitude estimates increased with increased
stimulus speed, but there was little indication of an
effect of stimulus pattern (disparity modulation type).
The latter nding is expected in the monocular
conditions because the disparity modulation would not
be visible without stereopsis. For the binocular
conditions it suggests that the particular pattern of
disparity-dened features was not critical for the
vection enhancement with binocular viewing.
A repeated-measures ANOVA (with Greenhouse-
Geisser correction where appropriate) indicated a
Figure 4. Vection magnitude ratings in Experiment 2 as a function of disparity modulation waveform, viewing condition (monocular or
binocular), and stimulus velocity. Each bar represents the mean of 15 subjects for the given condition; error bars indicate 61
standard error of the mean.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 11
signicant main effect of stimulus speed, with higher
speeds producing stronger vection ratings, F(1, 14)
30.35, p , 0.001, g
2
p
0.684. There was also a main
effect of viewing condition, F(1, 14) 4.75, p 0.047,
g
2
p
0.253, and an interaction between disparity
waveform type and viewing condition, F(2.22, 31.07)
4.512, p 0.016, g
2
p
0.244. The main effects of
disparity waveform and xation were not signicant
F(2.76, 38.69) 0.37, p 0.762, g
2
p
0.025 and F(1, 14)
0.45, p 0.513, g
2
p
0.031, respectivelynor were
there any other signicant interactions. One subject
reported much smaller vection magnitude under
monocular conditions than other subjects; the analyses
repeated with this subject removed yielded the same
pattern of results as with the full data set.
The interaction of disparity waveform and viewing
type was analyzed by looking at the simple main effects
of viewing for each waveform. This interaction was
expected since the waveform was dened by cyclopean
featureshence invisible monocularlyso if there was
an effect of waveform it should be apparent only under
binocular viewing. Binocular viewing increased vection
magnitudes, but this increase was signicant only for the
sine and square waveforms. Binocular viewing on
average generated 8.2 (63.49 SEM) and 9.3 (63.49)
increases in vection magnitude ratings, respectively
F(1, 14) 5.93, p0.033, g
2
p
0.285 and F(1, 14) 7.11,
p 0.018, g
2
p
0.337, respectivelywith smaller effects
for the sawtooth and triangle waveformsmean rating
increases of 7.1 (63.79) and 4.6 (63.00), respectively;
F(1, 14) 3.50, p0.083, g
2
p
0.200 and F(1, 14) 2.34,
p0.148, g
2
p
0.143, respectively. Thus, while binocular
viewing generally increased vection magnitudes, the
effect depended on the type of disparity waveform.
Vection time course data
An equivalent repeated-measures ANOVA on vec-
tion onset indicated only a signicant effect of stimulus
speed, F(1, 14) 21.01, p , 0.001, g
2
p
0.600. Vection
latency was shorter for the 0.163 m/s condition (5.99 6
1.42 s) than for the 0.082 m/s condition (8.37 6 1.85 s)
when averaged across the other variables and observ-
ers. Unlike for the magnitude measure, the effect of
viewing condition did not reach signicance: Mean
vection latency was lower for binocular compared with
monocular viewing by 0.83 6 0.45 s, F(1, 14) 3.43, p
0.085, g
2
p
0.197, and all other main effects and
interactions were not signicant.
Discussion
The nding that binocular viewing of RDS stimuli
produces stronger vertical vection than monocular
viewing is consistent with the hypothesis that move-
ment of stereoscopically dened form contributes
generally to the perception of linear self-motion.
However, this improvement was modest and depended
on the type of disparity waveform. It is possible that the
modest effects were due to vection induced by
monocular stimulation already being relatively strong
and saturated, leaving little room for stereoscopic form
to have an effect (i.e., ceiling effects).
It is also possible that the vection increases found for
binocular viewing were not due to the presence of
stereoscopic features but rather to the fact that two eyes
were being stimulated as opposed to only one.
However, differences in binocular and monocular eld
of view cannot explain our current results, as the
stimulus was contained well within the visual eld of
both eyes. On the other hand, binocular inputs are
known to sum in some cases to produce a stronger
signal-to-noise ratio. Hess et al. (2007) argued that
global motion processing occurs after the site of
binocular combination and that binocular improve-
ments in their global coherence detection tasks could be
attributed to binocular contrast enhancement. While
binocular summation may play a role in the binocular
advantage found in this experiment, it cannot be the
whole story because binocular summation cannot
explain the effect of disparity waveform typea
stimulus feature that is available only at a cyclopean
level of processing. The next experiment addressed
these issues of monocular self-motion signal strength
and binocular summation.
Experiment 3: Short-lifetime
cyclopean stimuli
The third experiment compared vection elicited by
stimuli with and without cyclopean features. This
allowed us to dissociate the effects of stereoscopically
dened features from binocularity. We used RDS stimuli
with limited dot lifetime to weaken the monocular dot
motion signals driving vection so that the stereoscopic
contribution could be assessed more sensitively. By using
short-lifetime dots, the monocular motion signals were
degraded and less likely to dominate and saturate the
vection response, allowing us to probe for the effects of
moving cyclopean form on vection.
Methods
A total of 13 nave observers (six males and seven
females ranging in age from 19 to 49 years; mean age 28
6 8.1 years) participated in Experiment 3.
Trials were randomized and grouped into blocks of
eight trials, with the standard stimulus presented before
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 12
each block. A total of 16 of these blocks were run over
two sessions for each subject. The standard stimulus
was a short-lifetime (dot lifetime of 10 frames) RDS
depicting a sinusoidal disparity wave moving upward at
0.082 m/s with xation on a central xation cross. The
independent variables were (a) the type of disparity
waveform (square wave or sinusoid; note that when
disparity amplitude was zerosee belowa at frontal
surface was dened), (b) stimulus speed (0.082 or 0.163
m/s), (c) xation (xation or free viewing), (d) disparity
direction, (e) disparity amplitude (5.44- or 0-arcmin
peak), and (f) dot lifetime (5 or 10 frames).
Results
Likelihood of vection induction
The main independent variables of interest in this
experiment were (a) disparity amplitude, which deter-
mined whether moving disparity-dened three-dimen-
sional surface features were present, and (b) dot
lifetime, which was intended to modulate the strength
of monocular motion signals. Subjects were less likely
to experience vection on ve-frame lifetime trials (74%)
than on 10-frame lifetime trials (92%). A logistic
regression of the dichotomous variable of vection
presence on the independent variables indicated that
likelihood of vection was inuenced by dot lifetime
(Wald v
2
94.84, p , 0.001), speed (Wald v
2
18.54, p
, 0.001), disparity (Wald v
2
29.88, p , 0.001), and
xation (Wald v
2
74.09, p ,0.001). While dot lifetime
had the greatest inuence, vection was also more likely
to occur for faster (compared with slower) stimulus
speeds, with xation (compared with free viewing), and
with disparity-dened depth corrugations (compared
with a disparity-dened at frontal plane).
Vection strength rating data
Figure 5 shows mean vection magnitude ratings as a
function of dot lifetime for each combination of speed
and disparity amplitude. Vection strength ratings were
larger for the depth corrugation (disparate) than for the
at (zero disparity) conditions for all combinations of
speed and dot lifetime. For the slower speed case, the
enhancement from disparity-dened three-dimensional
surface features was more pronounced for the shorter,
ve-frame lifetime than for the 10-frame dot lifetime.
This is presumably the condition with the weakest
monocular motion signals, and the effect of disparity
modulation is clearest here. Figure 6 shows that the
enhancement effect of disparity-dened three-dimen-
sional surface features was greater for the sinusoidal
disparity modulation than for the square wave dispar-
ity modulation.
A repeated-measures factorial ANOVA on the
vection magnitude data revealed several interactions.
Vection strength ratings tended to increase with
disparity amplitude, dot lifetime, and stimulus speed
(Figure 5), and there was a three-way interaction
between these variables, F(1, 12) 5.23, p 0.040, g
2
p
0.305. Further analysis demonstrated that the
simple main effects of stimulus speed, disparity, and
lifetime held at all levels of each of the other two
variables: Marginal main effects were signicant for
lifetime, F(1, 12) 25.12, p , 0.001, g
2
p
0.677;
disparity, F(1, 12) 18.51, p 0.001, g
2
p
0.607; and
speed, F(1, 12) 21.36, p 0.001, g
2
p
0.640.
Marginal to this three-way interaction was a two-way
interaction between speed and lifetime, F(1, 12)
11.43, p 0.005, g
2
p
0.488, with this interaction
depending on disparity. Specically, the increase in
vection ratings with longer dot lifetimes was smaller
for the 5.44-arcmin disparity, slow-speed condition
(mean increase 10.3 6 3.73 SEM) compared with the
0-arcmin conditions at the slower speed (19.4 6 4.28)
or with the 0-arcmin and 5.44-arcmin conditions at
the higher speed (mean increases 22.5 6 4.10 and 20.9
6 4.36, respectively).
There was also an interaction between disparity
waveform and disparity amplitude, F(1, 12) 6.32, p
0.027, g
2
p
0.345 (see Figure 6). This interaction was
expected and reects the fact that when disparity
Figure 5. Vection magnitude ratings in Experiment 3 as a
function of dot lifetime and disparity amplitude (flat versus
corrugated). The circular symbols show data for a stimulus
speed of 0.082 m/s, and the square symbols show data for a
stimulus speed of 0.163 m/s. Filled symbols show data for 5.44-
arcmin disparity (corrugated), and open symbols show data for
0-arcmin disparity (flat). Mean data for 13 observers; error bars
indicate 61 standard error of the mean.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 13
amplitude was zero the sinusoidal and square wave
stimuli were identical (both at RDS). Thus, as
expected, there was no difference between the vection
strength ratings for the two waveforms when disparity
was zero (mean difference 0.66 60.685, p0.351, g
2
p
0.073), but when there was disparity, the sinusoid
produced signicantly more vection than did the square
wave case (3.95 6 1.67, p 0.035, g
2
p
0.312). There
was a signicant effect of disparity for both waveforms,
with more vection for disparate stimuli compared with
at stimuli: main effect of disparity, F(1, 12) 18.51, p
0.001, g
2
p
0.607. Signicant simple main effects of
disparity were also found for both sinusoid and square
wave stimuli.
There was also a signicant effect of xation type,
F(1, 12) 6.92, p 0.022, g
2
p
0.366, with vection
magnitude increased for xation compared with free
viewing.
Vection time course data
Vection onset times tended to decrease with
increasing dot lifetime, stationary xation, disparity
amplitude, and stimulus speed (Figure 7). Consistent
with the magnitude estimation results, latency mea-
sures indicated that vection was enhanced when
disparity-dened features were present compared with
when they were not (corrugated versus at condi-
tions). For the at, free-view case, vection was
typically not obtained when dot lifetime was ve
frames (median onset latency was the length of the
trial). Latency in the at condition was more
comparable with the disparity corrugation condition
in the 10-frame, free-view case and for both lifetimes
with xation. Nevertheless, in all cases median vection
latency was shorter in the disparate conditions
compared with the zero-disparity conditions. Fixation
had a large effect on vection latencies, which were
shorter under xation than free-view conditions. This
xation enhancement was smallest for the strongest
vection condition (10 frame, 0.163 m/s), probably
reecting a oor effect because vection latencies were
typically short and near those found for the strong
vection in the RDS conditions in Experiment 1.
Censored regression indicated that there were
signicant main effects of lifetime, t(12) 7.87, p ,
0.001, d 4.55; disparity, t(12) 6.11, p , 0.001, d
3.53; speed, t(12) 2.389, p 0.034, d 1.38; and
xation, t(12) 7.08, p , 0.001, d 4.09. There was
also a two-way interaction between lifetime and
disparity, t(12) 4.54, p , 0.001, d 2.62, with the
effect of disparity signicant for both lifetimes but
smaller at the longer lifetime compared with the shorter
lifetime (mean difference between disparity levels of
1.23 6 0.529 and 4.86 6 1.30 s, respectively). A two-
way interaction between xation and lifetime was also
signicant, t(12) 3.14, p 0.008, d 1.81, and
reected a larger decrease in vection latency with
xation for the shorter-lifetime dots compared with the
longer-lifetime dots.
Discussion
Our use of short-lifetime dots appeared to have the
desired effect of weakening the monocular vection
Figure 6 . Waveformdisparity interaction in vection magnitude ratings in Experiment 3. Mean data for 13 observers collapsed across
velocity and fixation conditions; error bars indicate 61 standard error of the mean.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 14
stimulus. In Experiment 2, vection was reported on
98.8% of the trials versus only 83.3% of trials in the
current experiment. Similarly, vection onset was
delayed relative to Experiment 2, suggesting a weaker
vection stimulus in the current experiment (12.65 6
2.20 s vs. 7.18 6 1.63 s averaged across all trials). Note
that vection magnitude is a relative measure dened by
the standard and thus cannot be used to compare
vection strength across the two experiments. The
weakened monocular stimulus used in this experiment
appears to have made the experiment more sensitive to
effects of stimulus parameters, and we found signicant
effects of xation type as well as interactions that were
not observed in the previous experiments.
Figure 7. Onset latency in Experiment 3 as a function of dot lifetime, stimulus speed, disparity, and fixation. Solid lines indicate
conditions with no disparity modulation (flat), and dashed lines indicate disparity-modulated patterns (corrugated). Median data are
shown for each condition; error bars indicate 95% confidence intervals.
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 15
General discussion
Binocular contributions to vection
These experiments identied a clear binocular
contribution to the production of vertical linear vection
from lamellar optic ow. While purely cyclopean
motion did not generate compelling vection on its own
(Experiment 1), binocular viewing of stereoscopic
(three-dimensional) optic ow produced more compel-
ling vection than either the monocular viewing of these
same optic ow displays or the binocular viewing of
nonstereoscopic (at) displays (Experiments 2 and 3).
Importantly, the addition of cyclopean three-dimen-
sional surface features to binocularly viewed optic ow
displays was found to signicantly enhance vection.
This stereoscopic enhancement was more apparent
when the monocular motion signals were weakened (by
reducing RDS dot lifetimes; Experiment 3).
As we outlined in the Introduction, there are several
possible ways that binocular motion stimulation could
have contributed to vection processing. While increased
eld of view and binocular summation are probably
important factors in many natural situations, they
cannot explain the binocular enhancements found here
because the stimuli were always constrained to the
binocular visual eld and because enhancements were
found for stereoscopic compared with equivalent
binocular but nonstereoscopic displays. Thus, the
present experiments suggest a clear contribution of
depth from binocular stereopsis to the experience of
vection.
Importantly, the current ndings demonstrate that
binocular vection enhancements are not restricted to
conditions where changing-disparities and interocular-
velocity differences provide extra information about
motion in depth (e.g., those examined previously by
Palmisano, 1996, 2002). Since our displays all simulated
self-motion parallel to a frontal cyclopean surface,
these ndings reveal a new binocular contribution to
vection enhancement.
Mechanisms underlying these binocular
enhancements
Improvements in perceived rigidity of the scene
(Nakamura, 2010) due to stereopsis are also not likely
to explain the binocular advantage found in the present
study. While stereoscopic depth and motion parallax
could constrain and disambiguate each other in a rigid
scene (Di Luca, Domini, & Caudek, 2007; Richards,
1985), this is not the case with the stereoscopic displays
in the present experiments. In our displays with
stereoscopic depth, the lack of motion parallax between
near and far parts of the display should have conicted
with stereoscopic depth and, if anything, degraded the
perceived rigidity of the scene. For instance, in
Experiment 3 all the dots in a given display drifted
vertically at the same speed. Thus, the monocularly
available motion cues were consistent with a frontal
plane drifting vertically and inconsistent with a rigid
interpretation of the cyclopean corrugated surface. If
rigidity were the driving force then we would expect
that vection would have been stronger for the at, zero-
disparity stimulus than for the corrugated surface, but
in fact the opposite was true.
Similarly, the binocular vection enhancement was
not likely due to improved perception of layout or to
segregation of gure from ground. While binocular
viewing would have provided disparity cues to segre-
gate the near physical aperture from the more distant
motion display, this cue was available in all conditions
with binocular viewing regardless of the disparity in the
random-dot stimulus. Since the aperture was always
stereoscopically segregated from the moving stimulus
on the television monitor in Experiment 3 (by real, as
opposed to simulated, binocular disparities), this
cannot explain the cyclopean vection advantages we
found.
1
Similarly, binocular cues to environmental
layout were not likely to play a determining role either
since, as noted above in the discussion of rigidity, the
disparity-specied layout and the motion-dened lay-
out were not consistent. Furthermore, the theoretical
stereoscopic depth between peaks and valleys was 23
cm, which, although appreciable, was much smaller
than the viewing distance or the separation between the
aperture and screen. As a result, the stereoscopic depth
modulations would have provided relatively little
additional information about environmental layout.
Thus, the most parsimonious explanation of our
results is that the stereoscopic displays provided
cyclopean features that moved and were interpreted by
the visual system as the consequence of self-motion.
These features enhanced the motion of the dots
themselves and produced a more compelling vection
stimulus.
Self-motion from higher-order motion stimuli
The binocular stimuli in Experiments 1 and 2 and the
disparate stimuli in Experiment 3 all provided moving
stereoscopic three-dimensional surface features. In
persistent or short-lifetime RDS stimuli these moving
surface features provide additional motion signals that
could reinforce and strengthen the monocularly visible
optic ow of the dots. It appears that the visual system
treated these cyclopean features as part of the static
environment and attributed their motion to self-
motion. To our knowledge this is the rst demonstra-
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 16
tion that stereoscopic cyclopean stimuli can drive linear
vection, although there is other evidence of cyclopean
contributions.
For instance, Wolfe and Held (1980) designed an
experiment to investigate binocular contributions to
circular vection using dichoptic apparent motion. They
illuminated a rotating optokinetic drum with separate
strobe lights for the left and right eyes. When the
ashes were synchronized and in phase, vection was
minimal for ash rates below 7 Hz, after which vection
magnitude increased with strobe frequency. When the
ashes were out of phase in the two eyes, vection
magnitudes were larger than for in-phase ashes at the
same rate, consistent with the observers combining the
two eyes input to form a signal with a higher effective
ash rate. The temporal offset between ashes would
also produce disparity signals because of the shift of the
stimulus during the interash interval. However, the
facts that the resulting disparity would be ambiguous
as both the preceding and following frames are
potential matchesand that Wolfe and Held found
similar results for stereoblind individuals suggest that
this binocular vection enhancement was not mediated
by stereopsis.
The studies by Palmisano (1996, 2002) discussed in
the Introduction suggested that stereoscopic motion in
depth could provide a stimulus to vection in depth. We
found that vection from lamellar ow can also be
enhanced by stereoscopic features moving in the frontal
plane. This suggests that stereoscopic enhancement
could be a more general response to moving cyclopean
features and not specic to stereoscopic motion in
depth. Lowther and Ware (1996) also found that both
circular vection and linear vection onset latencies were
lower for stereoscopically presented stimuli compared
with nonstereoscopic binocular displays, but they did
not explore the nature of the stereoscopic benet.
Motion of cyclopean features provides an alternative
vection stimulus to the movement of luminance-dened
features (rst-order motion). This is consistent with
other evidence that vection can be elicited by motion
stimuli that are not luminance dened. So-called
second-order (and third-order) motion refers to motion
percepts elicited by stimuli such as contrast envelope,
texture, and icker that can dene moving features
without corresponding moving luminance-dened fea-
tures. Gurnsey, Fleet, and Potechin (1998) presented
displays simulating forward motion through a tunnel
and varied the relative contribution of rst- and
second-order motion in the stimulus. Both motion
aftereffect and vection durations were strongest when
the stimulus contained coherent rst-order motion
energy. Similarly, Seno and Palmisano (2012) found
that forward vection was enhanced by addition of
vertical oscillation of the simulated viewpoint when the
oscillatory movement was dened by rst-order motion
but not second-order motion. These ndings are similar
to the results of our Experiment 1, where vection
responses were small or absent in the absence of rst-
order motion of the dots (i.e., in DRDS displays
compared with RDS displays). Consistent with a lack
of an inuence of DRDS stimuli on self-motion,
presenting a visual display as a DRDS reportedly
eliminates the stabilizing effect of vision on sway
(Kelly, Riecxe, Loomis, & Beall, 2008).
In Gurnsey et al.s (1998) second experiment they
modulated the relative contribution of rst-order
motion by varying the scale of the noise carrier. While
motion aftereffect duration decreased as the relative
amount of rst-order motion energy declined, they
found that duration of vection was unaffected. They
concluded that vection was driven by optic ow that
combines both rst- and second-order motion energy.
Analogously, in Experiment 3 we degraded the
effectiveness of the rst-order motion signal by limiting
dot lifetime and demonstrated a contribution of
cyclopean motion signals when combined with weak
but consistent rst-order dot motion.
In many natural situations the self-motion signals
from cyclopean features, second-order features, and
luminance-dened features are consistent and redun-
dant. In the present experiments the cyclopean and
luminance-dened motions were either consistent or
effectively absent. It will also be of interest in future
work to examine how these cues combine when either
weakly or strongly inconsistent. In a pilot experiment
we designed a stimulus consisting of a moving
cyclopean sinusoidal grating but with persistent rather
than short-lifetime (scintillating) dots. Each dot in these
displays maintained its cyclopean position; that is, its
disparity changed as the disparity wave travelled over
it. This display is similar to the DRDS case where
monocular dot motion is on average zero; however, in
these persistent displays the dot motion was unambig-
uously zero. As such the conict with the monocular
motion cue is stronger than the DRDS case (Allison &
Howard, 2000). Consistent with our DRDS results,
there was no vection reported by four out of the ve
observers in this pilot experiment; the other reported no
vection on most trials but weak and nonspecic self-
motion (instability) on a few trials.
Effects of cyclopean pattern and phase
This study examined for the rst time the vection
and vection enhancement provided by cyclopean
motion. When examining the motion of purely
cyclopean features, the disparity modulation of the
surface is (in principle) an important consideration.
Accordingly, a number of different disparity modula-
tions (sawtooth, sinusoidal, square wave, and triangle
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 17
wave) were examined to see whether the type of display
modulation mattered.
The type of disparity modulation is a feature that by
denition exists in the cyclopean domain. Not surpris-
ingly, then, we found no effect of the disparity
waveform pattern or phase (disparity sign) on vection
in Experiment 1, consistent with the DRDS producing
no vection. In contrast, the disparity modulation
waveform had modest interaction effects with view type
(binocular versus monocular) in Experiment 2 and
waveform disparity amplitude (5 vs. 0 arcmin) in
Experiment 3. These interactions were expected as the
disparity waveform had no meaning for monocular or
zero-disparity stimuli.
The disparity waveform was of interest in these
experiments for two reasons. First, the cyclopean
motion is dened only where the disparity changes since
the motion stimulus does not exist except at cyclopean
features. This is potentially important because vection
depends on the number and density of moving features
(Brandt, Wist, & Dichgans, 1975). The sinusoid has
continually varying disparity; therefore, cyclopean
motion is dened throughout the stimulus while it is
dened only at the discontinuities for the square wave.
Consistent with this, we found that signicantly more
vection magnitude was produced by the sinusoidal
stimulus compared with the square wave stimulus in
Experiment 3. However, the difference was modest, and
in Experiment 2 the binocular improvement was similar
for the sine and square wave patterns. These ndings
suggest that the continuity of moving cyclopean
features was only a minor factor in the production of
vection enhancements, particularly in the presence of
strong monocular motion as in Experiment 2.
Second, spatial and temporal frequency changes are
confounded in drifting sinusoidal gratings (Kohly &
Regan, 1999) and, to a lesser extent, in other periodic
patterns. Variation in harmonic content and phase in
different patterns can help to alleviate this confound.
Although the binocular improvement in Experiment 2
was signicant only for the sinusoid and square wave
cases, the improvements were only slightly smaller with
the sawtooth wave: These three patterns have different
spatiotemporal spectra but produced similar binocular
improvements. Thus, covariation of spatial and tem-
poral frequency does not appear to be a signicant
factor in the current experiment. Similarly, the dispar-
ity sign affects the phase of the waveform and hence the
features appearing near xation at the start of the trial.
Furthermore, the sawtooth wave presents only one
direction of surface slant that is determined by the sign
of the disparity, and there are known anisotropies and
biases in the perception of stereoscopic slant about a
horizontal axis (e.g., Allison, Howard, Rogers, &
Bridge, 1998). However, we did not nd effects of the
disparity sign and thus there was no evidence that these
phase-dependent differences affected vection.
The smallest binocular improvement in Experiment
2 was observed for the triangle wave. Except at the
peaks, this stimulus has only gradients of disparity
(alternately slanted sections). Observers are relatively
insensitive to stereoscopic slant, particularly when
presented in these so-called hinge arrangements (Gil-
lam, Blackburn, & Brooks, 2007; Gillam, Chambers, &
Russo, 1988). This insensitivity has been conrmed for
both motion and stereopsis (Allison, Rogers, &
Bradshaw, 2003) and could explain why the binocular
improvement was somewhat smaller for the triangle
wave modulation compared with the other waveforms.
Effects of fixation
The role that stationary xation plays in the
induction of (typically nonstereoscopic) vection has
long been a matter of some debate. Some previous
work has suggested that circular vection tends to be
weaker when observers follow the visual-inducing
pattern compared with static xation conditions (de
Graaf, Wertheim, & Bles, 1991; Fushiki, Takata, &
Watanabe, 2000). This increase in circular vection with
xation has not always been found; for example, de
Graaf et al. (1991) found the effect only when following
and xation conditions were presented sequentially in
such a way that they could be compared immediately.
Consistent with an earlier study by Dichgans and
Brandt (1978), they found no effect of xation when the
conditions were presented separately (i.e., in isolation).
Similarly, while vection in a physically tumbling room
was found to increase or decrease with xation for
individual subjects, there was no consistent overall
effect (Allison, Howard, & Zacher, 1999). Central
xation on a stationary target reportedly improves
vection for centrally presented, but not peripherally
presented, horizontal linear vection stimuli compared
with free viewing (Tarita-Nistor, Gonz alez, Spigelman,
& Steinbach, 2006), although alternating gaze between
central and peripheral locations increased looming
vection compared with stable central xation (Palm-
isano & Kim, 2009).
In the present studies we did not nd a signicant
effect of xation on ratings of stereoscopic (and
nonstereoscopic) vection strength in the rst two
experiments. However, vection onset latencies were
slightly reduced with xation compared with free
viewing in Experiment 1. Furthermore, in Experiment
3, with brief-lifetime dots we found that vection
magnitudes were increased and onset latencies were
reduced with xation compared with free viewing.
Monocular motion cues were diminished by the use of
short-lifetime dots in this experiment, and it may be
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 18
that this allowed a more subtle effect of xation to be
measured. Contrary to this proposal, Tarita-Nistor et
al. (2006) found that the enhancement of vection with
xation was more pronounced for their larger, more
compelling inducing stimuli.
We found that when xation had an inuence,
vection latency was reduced and magnitude increased
by the presence of the stationary xation point.
However, the xation manipulation introduced both a
feature to the display (the xation cross) and instruc-
tions to xate. This is also true of most previous studies
looking at the effects of xation. Previous research has
assumed that the act of xation is the important
manipulation and have tried to explain the increase in
vection sensation through mechanisms such as the
Aubert-Fleischl phenomenon, where the perceived
speed of a moving target is higher during steady
xation compared with when it is tracked (de Graaf et
al., 1991).
However, there could also be more direct effects of
the xation point on vection. First, addition of a
xation point introduced an additional reference for
relative motion, which has been reported to increase
vection (Howard & Howard, 1994). Furthermore, in
our study there was also a very strong percept of
induced motion apparent in the xation point.
Researchers have distinguished between vection-en-
trained induced motion, where objects that appear
xed with respect to the head appear to move with it
during vection, and egocentric induced motion, where
a moving stimulus induces opposite motion of another
stimulus without the percept of self-motion (e.g.,
Heckmann & Howard, 1991). In principle, the xation
point in our experiments could have been subject to
both these effects. Thus, while there was a large effect
of xation on vection in Experiment 3, it is possible
that this effect could have been mediated by induced
motion rather than by xation per se (although
xation plays a key role in many theories of induced
motion). We are currently exploring the role of
xation stability versus induced motion in the
production of vection.
Relation of the current findings to
neurophysiology
Much of the visual cortex is sensitive to motion or
disparity, and thus the neurophysiological underpin-
nings of the effects of cyclopean motion on vection are
difcult to localize precisely. Cyclopean form depends
on binocular combination and disparity sensitivity, but
substantial numbers of cells responsive to these features
and to luminance-dened motion are found as early as
V1 (Hubel & Wiesel, 1962; Poggio & Fischer, 1977).
However, disparity-sensitive cells in V1 do not signal
relative disparity or cyclopean form, and the earliest
evidence for representations of cyclopean edges is in V2
(Bredfeldt & Cumming, 2006; Zhou, Friedman, & von
der Heydt, 2000). Representations of cyclopean shape
appear to arise later in the ventral pathway, and cells
sensitive to disparity-dened form have been reported
in the inferior temporal cortex (Tanaka, Uka, Yosh-
iyama, Kato, & Fujita, 2001).
One possibility is that such cells produce local
cyclopean features and related motion signals and that
these motion signals then feed into global self-motion
processing (see Patterson, 1999 for a comprehensive
review of this proposal). Disparity sensitivity is a
common feature throughout the motion-sensitive
visual pathway. Thus, these signals would be pro-
cessed much as luminance-dened motion features
arising in V1 are processed and integrated in
subsequent stages of the motion-sensitive pathways
such as the medial temporal (MT) and medial superior
temporal (MST) areas. Consistent with this, De-
Angelis, Cumming, and Newsome (1998) reported
that stimulating clusters of MT neurons in rhesus
monkeys produced predictable bias in coarse depth
judgments. Area MT is intimately linked with primate
motion processing and is selective for disparity, which
could help segregate motion signals at different
distances (Bradley, Qian, & Andersen, 1995). One
problem with the proposal that cyclopean motion
features are treated identically to luminance-dened
motion is that disparity signals in MT do not appear
to code for disparity-dened boundaries or cyclopean
motion and may be better suited to guiding distance-
dependent motor action. More generally, it has been
proposed that disparity processing in ventral cortical
areas may support representations of three-dimen-
sional shape while dorsal cortical areas support motor
action and segregation of motion signals (for review
see Parker, 2007). To the extent that this is accurate,
contributions of moving cyclopean form must rely
either on communication between the dorsal and
ventral streams or on some unknown representation of
stereoscopic shape in the dorsal stream.
The contribution of moving cyclopean stimuli to
vection may be more specic to the processing of self-
motion and the perception of vection rather than
simply providing an alternative input. For instance,
cortical area MST, which has been associated with
egomotion processing, is sensitive to both patterns of
optic ow and binocular disparity (Roy, Komatsu, &
Wurtz, 1992). Some cells are jointly tuned to disparity
and motion and some are sensitive to depth from
relative disparity and motion parallax (Eifuku &
Wurtz, 1999; Upadhyay, Page, & Duffy, 2000).
Cardin and Smith (2011, p. 1246) proposed that a
signature of a brain region encoding egomotion is
that it responds well to optic ow and the response
Journal of Vision (2014) 14(12):5, 123 Allison, Ash, & Palmisano 19
is systematically enhanced when depth cues are
consistent with observer motion. The only region
they found with these properties was V6, and they
suggested that this area was important for integrating
stereopsis and motion to dissociate egomotion from
object motion. They found that when stereoscopic
layout was consistent with the optic ow stimulus
there was an enhancement of the response to optic
ow. This facilitation of optic ow by consistent
stereopsis appears similar to our nding that consis-
tent cyclopean motion enhanced vection from mon-
ocular optic ow even when it was ineffective on its
own. However, the stimuli used by Cardin and Smith
(2011) were not cyclopean and provided layout cues
as well as cyclopean form. Thus, any specic links
between specialized processing of cyclopean form and
egomotion-specic cortical regions remain specula-
tive.
Conclusions
Previous work has demonstrated an enhancement
of vection with binocular stereopsis (Palmisano, 1996,
2002). We outlined possible factors underlying this
binocular enhancement and isolated one important
component: that of the motion of stereoscopically
dened features. In sum, the current experiments
provide compelling evidence for the inuence of
cyclopean features but only when paired with consis-
tent monocular self-motion signals. Vection from
purely cyclopean stimuli (DRDS) was not obtained
despite robust perceptions of the motion of their
cyclopean features. However, the DRDS stimuli
depicting translating gratings have lower apparent
contrast than do RDS stimuli, which might in turn
have resulted in lower apparent velocities in the
DRDS case (Brooks, 2001; Thompson, 1982). Such
DRDS stimuli also have motion discrimination
thresholds that are slightly (Kohly & Regan, 1999;
Portfors & Regan, 1997) or considerably (Harris &
Watamaniuk, 1996) higher than for equivalent lumi-
nance gratings. Also, the scale of features in the
cyclopean gratings was coarser than that of the
monocular features, which could have possibly pro-
duced spatial frequency-dependent apparent speed
reductions in the cyclopean gratings (Diener, Wist,
Dichgans, & Brandt, 1976). Thus, while we have
compelling evidence that cyclopean features can
promote self-motion perception when combined with
monocular motion signals, more work is necessary
before we can convincingly conclude that cyclopean
motion cannot produce vection on its own.
Keywords: vection, cyclopean vision, self-motion,
lamellar ow, stereopsis
Acknowledgments
A brief report on this work was published as an
abstract and presented at the Asia-Pacic Conference
on Vision 2013. This research was supported by grants
from the Ontario Media Development Corporation
and NSERC (Canada) as well as by an Endeavour
Fellowship to A. A. from the government of Australia.
Commercial relationships: none.
Corresponding author: Robert S. Allison.
Email: allison@[Link].
Address: Department of Electrical Engineering and
Computer Science, York University, Toronto, Canada.
Footnote
1
Stereoscopic segregation of the aperture from the
random-dot display might have been a factor in the
binocular enhancement of vection (compared with
monocular viewing conditions) in Experiment 2.
However, in Experiments 1 and 3, all conditions were
binocular.
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