Journal of Vision (2024) 24(2):4, 1–14
Effects of binocular disparity on binocular luminance
combination
Goro Maehara*
Yiqian Wang*
Ikuya Murakami
This study aimed to examine the effects of binocular
disparity on binocular combination of brightness
information coming from luminance increments and
decrements. The point of subjective equality was
determined by asking the observers to judge which
stimulus appeared brighter—a bar stimulus with
variable disparity or another stimulus with zero
disparity. For the bar stimulus, the interocular
luminance ratio was varied to trace an equal brightness
curve. Binocular disparity had no effect on luminance
increments presented on a gray or black background. In
contrast, when luminance decrements were presented
on a gray background, non-zero disparities elevated
points of subjective equality for stimuli with interocular
luminance differences. This means that the binocular
brightness combination of the two monocular signals
shifted from winner-take-all summation toward linear
averaging. It has been argued that this effect may be
caused by non-zero binocular disparities attenuating
interocular suppression, which is deemed to operate
normally with zero disparity.
Introduction
Researchers have examined how visual signals for
brightness delivered to the two eyes are combined
to yield brightness perception in binocular viewing,
hereafter referred to as binocular brightness. In their
review of previous studies, Blake and Fox (1973) argued
that, although viewing with both eyes open is not
dramatically different from viewing with only one eye,
sensitive experiments can reveal differences between
binocular and monocular brightness. This trend dates
back to the work of Fechner (1860), who noted that
the binocular brightness of the light spot decreases
as the luminance in one eye increases from zero. This
Citation: Maehara, G., Wang, Y., & Murakami, I. (2024). Effects of binocular disparity on binocular luminance combination. Journal
of Vision, 24(2):4, 1–14, https://doi.org/10.1167/jov.24.2.4.
https://doi.org/10.1167/jov.24.2.4 Received June 7, 2023; published February 20, 2024
This work is
Downloaded from jov.arvojournals.org onlicensed under a Creative Commons Attribution-NonCommercial-NoDerivatives 4.0 International License.
03/23/2024
1
Department of Human Science,
Kanagawa University, Kanagawa, Japan
Department of Psychology,
The University of Tokyo, Tokyo, Japan
Department of Psychology,
The University of Tokyo, Tokyo, Japan
phenomenon is referred to as Fechner’s paradox.
Levelt (1965) and Engel (1970) conducted controlled
experiments on binocular brightness combinations.
They found that, when both eyes were sufficiently
illuminated, binocular brightness appeared to be
perceptually matched among the conditions in which
the linear averages of the two monocular luminance
values were equivalent. In contrast, when the luminance
in one eye was much lower than that in the other eye,
the eye with higher luminance determined the binocular
brightness (winner-take-all), which is consistent with
Fechner’s paradox.
The way of binocular brightness combination
is known to vary with the luminance polarity and
the luminance difference between a stimulus and its
background. According to Anstis and Ho (1998), when
a luminance decrement (a dark spot) was presented
on a white background, the binocular brightness
combination took the form of winner-take-all. That
is, a greater decrement in one eye always determined
the binocular brightness without binocular averaging.
Moreover, Baker, Wallis, Georgeson, and Meese (2012)
adopted a gray background, instead of black or white,
to test the binocular brightness of light and dark spots.
Their results showed that the equal brightness functions
for luminance increments deviated from the prediction
of linear averaging, being close to winner-take-all. The
functions for luminance decrements have also exhibited
compromised linearity for small decrements. Ding and
Levi (2017) reported similar results for a wider range of
target and background luminance. In Ding and Levi’s
(2017) model, interocular suppression is assumed to be
weak when the total luminance of the stimuli is low and
the luminance values are balanced between the two eyes,
resulting in linear averaging. If the total luminance is at
a medium level, the binocular brightness combination
is expected to show compromised linearity for both
ISSN 1534-7362 Copyright 2024 The Authors
 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami
luminance increments and decrements. This model was
recently expanded to explain depth perception (Ding &
Levi, 2021).
Interestingly, the binocular brightness of Ganzfeld
stimuli followed linear averaging over the entire
range of stimulus luminance, with no indication of
Fechner’s paradox (Bourassa & Rule, 1994). This
could be due to a weaker interocular suppression by
Ganzfeld stimuli than by spatially localized stimuli. In
contrast, binocular combinations have been reported
to shift from winner-take-all to linear averaging when
binocularly matched features are added to stimuli
in dichoptic color-saturation mixtures (Kingdom &
Libenson, 2015) and when the luminance contrasts of
surroundings are the same for both eyes in dichoptic
contrast mixtures (Wang, Ding, Levi, & Cooper, 2022).
This evidence suggests that suppression of the eye with
the lower contrast by the eye with the higher contrast
results in winner-take-all behavior. Binocularly matched
features would keep the interocular suppression
balanced between the two eyes, resulting in a shift of the
binocular combination toward linear averaging. This
notion is supported by studies showing that binocularly
matched features reduce the effects of dichoptic
masking (Jennings & Kingdom, 2019; Kingdom &
Wang, 2015).
The aforementioned studies focused on situations
in which binocular stimuli were presented at the
corresponding retinal positions. Recently, Maehara
and Murakami (2020) reported that suprathreshold
grating stimuli with non-zero binocular disparity
appeared to have a higher luminance contrast than
those with zero binocular disparity. As simultaneous
stimulations at corresponding retinal points are known
to elicit strong interocular suppression (Blake, 1989;
for an opposing view, see a dynamical model proposed
by Said & Heeger, 2013), the authors attributed
the perceptual enhancement of luminance contrast
to weaker interocular suppression in stimuli with
binocular disparities. In contrast, binocular summation
is assumed to be functional even for stimuli with
binocular disparity, because the same mechanisms
may underlie binocular summation and stereopsis
(Kingdom, Yared, Hibbard, & May, 2020; Lema &
Blake, 1977; May & Zhaoping, 2022; Rose, Blake, &
Halpern, 1988).
The present study aimed to examine the effects
of binocular disparity on the binocular brightness
combination of luminance increments and decrements.
We measured the point of subjective equality (PSE)
in brightness between a standard stimulus with zero
disparity and a comparison stimulus with a certain
interocular excursion ratio and a zero, crossed, or
uncrossed binocular disparity. If binocular disparity
caused weaker interocular suppression, then the
binocular brightness combination for a stimulus with
non-zero as opposed to zero disparity would be shifted
Downloaded from jov.arvojournals.org on 03/23/2024
2
away from winner-take-all toward averaging. There
were three conditions in the luminance excursion (i.e., a
generic term to indicate a difference, either incremental
or decremental, in the luminance values between the
stimulus and background) of the comparison stimulus:
a decrement on a gray background, an increment
on a gray background, and an increment on a black
background. We used a short vertical bar-shaped
stimulus as the comparison stimulus so that it did
not stimulate the corresponding retinal positions in
the two eyes when presented with non-zero binocular
disparity. To avoid a masking effect on the comparison
stimulus, the standard stimulus was a pair of relatively
large rectangles presented within a region outside
the location of the comparison stimulus. Binocular
disparities within the fusion limit were selected
because preliminary observers had difficulty judging
the brightness in double vision when the luminance
excursions differed between the eyes.
Methods
Observers
Four adults naïve to the purpose of the experiment
and two authors (observers 5 and 6) participated
in Experiment 1. Four different naïve adults and
the two authors participated in Experiment 2. All
observers had normal or corrected-to-normal visual
acuity with normal stereopsis and provided written
informed consent. The present study followed protocols
approved by the institutional ethics committee and was
in accordance with the tenets of the Declaration of
Helsinki.
Apparatus
In a dimly lit room, a ViSaGe MKII Stimulus
Generator (Cambridge Research Systems, Rochester,
UK) with a 14-bit gray-level resolution was used
to present the stimuli on a CRT video monitor
(RDF223H; Mitsubishi, Tokyo, Japan). The display
resolution was 800 × 600 pixels, with a refresh rate of
100 Hz. The observers viewed the screen using a mirror
stereoscope (Chuo Precision Industrial, Tokyo, Japan).
The presentation area subtended a 13° × 13° arc in each
eye. The viewing distance was 57 cm.
Stimuli
There were three excursion conditions: a decrement
on a gray background, an increment on a gray
background, and an increment on a black background.
 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 3

The gray and black backgrounds had luminance values Left Eye Right Eye
of 35.1 cd/m2 and 0.2 cd/m2 , respectively. A square
frame subtending 9.9° × 9.9° was presented to each (a)
eye to aid binocular fusion (Figure 1). A fixation dot
was presented at the center of the presentation area
with two flanking dots positioned 1.5° to the left and
right (Figure 1a). Nonius dots were also presented 1.5°
above and below the fixation dot to the left and right
eyes, respectively. The luminance of these dots was 21.0
cd/m2 higher than the background luminance.
The standard stimulus was a pair of rectangles each
subtending 9.9° × 1.0° (Figures 1b and 1c). The stimuli (b)
were presented 3.0° above and below the fixation dot.
In Experiment 1, their luminance was 14 cd/m2 lower
in the luminance decrement condition (Figure 1b) and
14 cd/m2 higher in the luminance increment conditions
(Figure 1c) than the background luminance. In
Experiment 2, the luminance of the standard stimulus
was 21 cd/m2 lower than the background luminance.
As a comparison stimulus, a vertical bar subtending
0.2° × 1.1° was presented at the center (Figures 1d
and 1e). In Experiment 1, the excursion levels, either (c)
decremental or incremental, of the comparison stimuli
were chosen as 8.8, 10, 11.4, 13.1, 15, 17.2, and
19.7 cd/m2 except that, for one of the observers, the
excursion range had to be shifted to 10 to 22.5 cd/m2
for the luminance increment on a gray background
after the first session. In Experiment 1, three interocular
excursion ratios were chosen for the comparison
stimulus. In the equal-ratio (L:R = 1:1) condition, the
images of the left and right eyes had the same excursion. (d)
In the dichoptic conditions (L:R = 1:0.5 or 0.5:1), the
excursion in one eye was half the “base” excursion value
(i.e., one of the above-described excursion levels). In
Experiment 2, the excursion levels of the comparison
stimuli were 13.2, 15.1, 17.2, 19.7, 22.5, 25.8, and
29.6 cd/m2 , and five interocular excursion ratios (1:1,
1:0.75, 1:0.5, 0.75:1, and 0.5:1) were selected for the
comparison stimuli.
The binocular disparity of the comparison stimuli (e)
had three levels: zero (0°), crossed (−0.2°), and
uncrossed (0.2°) disparities. In the zero-disparity
condition, monocular images were displayed at the
center. The images were horizontally shifted by 0.1° in
opposite directions for the left and right eyes to create
non-zero disparities.

Procedures
The observers pressed a key to start each trial when
the Nonius and fixation dots appeared to be aligned
along the vertical meridian. After a 400-ms blank, the
standard stimulus was presented for 200 ms. After
another 600-ms blank, the comparison stimulus was
presented for 200 ms. The fixation dot disappeared
100 ms before the onset of the comparison stimulus
Figure 1. Stimulus examples. (a) A fixation dot and Nonius dots.
(b) The standard stimulus for the luminance decrement on a
gray background. (c) The standard stimulus for the luminance
increment on a gray background. (d) The comparison stimulus
for the luminance decrement on a gray background. (e) The
comparison stimulus for the luminance increment on a black
background.

Downloaded from jov.arvojournals.org on 03/23/2024

 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 4

but the other dots remained. The duration of each

(a) Decrement on Gray BG stimulus presentation was too short to evoke vergence in
20 response to a disparate stimulus. The observers judged
Zero Cross. Uncross. which stimulus was darker in the luminance-decrement
condition or brighter in the luminance-increment
15 condition.
In Experiment 1, each experimental block had 189
trials, consisting of three repetitive trials for each
10 combination of three binocular disparities (zero,
L L L R R R crossed, and uncrossed), three interocular excursion
PSE of the Comparison Stimulus Matched to the Standard Stimulus (cd/m²)

ratios (1:1, 1:0.5, and 0.5:1), and seven excursion levels

5
(see above). The order of the trials was randomized
within a block. The observers rested after completing
0 a block and adapted to the background luminance
for 30 seconds before starting each block. During
1:1 1:0.5 0.5:1
this adaptation period, the standard stimulus of the
impending block was presented for 200 ms every 3
seconds. The observers completed four blocks in a
(b) Increment on Gray BG
20 session devoted to one of three excursion conditions
(a decrement on a gray background, an increment
on a gray background, and an increment on a black
15 background). Three sessions were conducted for each
excursion condition; thus, 6804 (189 × 4 × 3 × 3) trials
were conducted for each observer. The order of the
10 sessions was counterbalanced among the observers.
L L L R R R In Experiment 2, each experimental block had
210 trials consisting of two repetitive trials for each
5 combination of the three binocular disparities (zero,
crossed, and uncrossed), five interocular excursion
ratios (1:1, 1:0.75, 1:0.5, 0.75:1, and 0.5:1), and seven
0 excursion levels (see above). The observers completed
1:1 1:0.5 0.5:1 four blocks per session. Four sessions were conducted
for a luminance decrement on a gray background; thus,
3360 (210 × 4 × 4) trials were conducted for each
(c) Increment on Black BG observer. All other procedures were the same as those
20
in Experiment 1.
We determined the brightness PSEs and 95%
15 confidence intervals (CIs) of the comparison stimuli
that should be perceptual matches to the standard
stimuli by fitting the log-Quick function to the data
10 using the Palamedes toolbox (Kingdom & Prins,
2016). There were 252 trials for each fit (36 trials × 7
L L L R R R excursion levels) in Experiment 1 and 224 trials (32 ×
5 7 excursion levels) in Experiment 2. PSE was defined
as the excursion corresponding to a 50% probability

0
1:1 1:0.5 0.5:1
←
Interocular Excursion Ratio the standard stimulus that was fixed across all conditions. The
of the Comparison Stimulus larger excursions (base PSEs) are plotted in the dichoptic
conditions (1:0.5 and 0.5:1). The letters L and R in the panels
Figure 2. Mean brightness PSEs in three binocular disparity indicate the eye (left and right) that received the designated
conditions and three interocular excursion ratios. The ordinate excursion. The error bars indicate the 95% CIs of the mean PSEs.
indicates the excursion (i.e., the absolute value of (a) Results for the luminance decrement on a gray background.
increment/decrement from the background luminance) (b) Results for the luminance increment on a gray background.
required to establish the brightness match to the brightness of (c) Results for the luminance increment on a black background.
→

Downloaded from jov.arvojournals.org on 03/23/2024

 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami
of judging the comparison stimulus as having greater
excursion than the standard stimulus.
Results of Experiment 1
Comparisons of PSEs
Figure 2 shows the mean PSEs for the three binocular
disparities (zero, crossed, and uncrossed) and the
three interocular excursion ratios (L:R = 1:1, 1:0.5,
and 0.5:1) plotted separately for the three excursion
conditions. When the excursions were unequal between
the eyes, a larger excursion, hereafter referred to as the
base PSE, was plotted. When the luminance excursions
were equal (1:1) for both eyes, the PSE was expected to
be approximately 14 cd/m2 , which was the luminance
excursion of the standard stimulus. The actual results
were roughly consistent with the predictions. As
shown in Figure 2, the PSE for the equal ratio (1:1)
was approximately 12 to 13 cd/m2 . Although these
measurements were slightly lower than the standard
excursion, this was possibly because the comparison
stimulus was presented near the fovea, whereas the
standard stimulus was presented within the parafovea
(3.0° of eccentricity).
If the binocular brightness combination was
winner-take-all, then the lengths of the bars would
be the same regardless of the interocular excursion
ratio. However, Figure 2 suggests the contrary because
the bars are generally longer for unequal ratios
(1:0.5 and 0.5:1) than for the equal ratio (1:1). In
contrast, if the binocular brightness combination
was a linear average, then the length of the bars
would be 4/3 times longer for the unequal ratios
than for the equal ratio. This prediction was more
or less applicable to the luminance increment on a
black background (Figure 2c). On the other hand,
for the luminance decrement on a gray background
(Figure 2a), the bars in the unequal ratios (1:0.5 and
0.5:1) did not support this linear-average prediction.
The data in the luminance increment on a gray
background were somewhat intermediate (Figure 2b).
Similar trends were observed in the individual
data, as shown in Figure 3 for two representative
observers.
We subjected the base PSE values to a two-way
repeated-measures analysis of variance (ANOVA)
with the interocular excursion ratio (1:1, 1:0.5, or
0.5:1) and binocular disparity (zero, crossed, or
uncrossed) as factors. In addition to the two-way
ANOVAs, we conducted Dunnett’s multiple-
comparison tests for each interocular excursion ratio.
In this multiple comparison, the base PSE at zero
binocular disparity was considered as the control
condition because our main concern was the effect
Downloaded from jov.arvojournals.org on 03/23/2024
5
of binocular disparity on the binocular luminance
combination.
For the luminance decrement on a gray background
(Figure 2a), the main effect of interocular excursion
ratio was significant, F(2, 10) = 40.6, p < 0.001, ηp 2
= 0.890. Between the two bilaterally symmetrical
conditions for unequal ratios (1:0.5 and 0.5:1), the
base PSEs were expected to be comparable. Thus, the
significant main effect presumably reflects the result
that the base PSE was smaller for the equal (1:1) ratio
(12.3 cd/m2 ) than for the unequal ratios (14.6 and 13.7
cd/m2 for the 1:0.5 and 0.5:1 ratios, respectively). The
two-way ANOVA also revealed a significant main effect
of binocular disparity, F(2, 10) = 11.2, p = 0.003, ηp 2
= 0.691, and a significant interaction, F(4, 20) = 3.45,
p = 0.027, ηp 2 = 0.409. According to Dunnett’s test
to clarify the effect of binocular disparity, the crossed
disparity yielded a significant elevation in the base PSEs
in the dichoptic conditions (1:0.5 and 0.5:1) but not
in the equal ratio (1:1) condition (p > 0.05). The 95%
CIs of the PSE elevations ranged from 0.230 to 1.80
cd/m2 for the 1:0.5 ratio (p = 0.014) and from 0.320 to
1.81 cd/m2 for the 0.5:1 ratio (p = 0.008). However, the
uncrossed disparity showed no significant effect (p >
0.05).
For the luminance increment on a gray background
(Figure 2b), the two-way ANOVA yielded a significant
main effect of interocular excursion ratio, F(2, 10) =
88.7, p < 0.001, ηp 2 = 0.947, indicating that the base
PSE was smaller for the equal (1:1) ratio (12.9 cd/m2 )
than for the unequal ratios (16.6 and 15.7 cd/m2 for the
1:0.5 and 0.5:1 ratios, respectively). The main effect of
binocular disparity was not statistically significant (p
> 0.05). The interaction between the two factors was
significant, F(4, 20) = 4.81, p = 0.007, ηp 2 = 0.490.
However, in Dunnett’s multiple-comparison tests,
neither crossed nor uncrossed disparities produced
significant changes in the base PSEs from the PSE under
the control condition (zero disparity) in any of the
interocular excursion ratios (p > 0.05). The significant
interaction may reflect some ratio-dependent differences
between the crossed and uncrossed disparities, the
origin of which could not be identified.
For the luminance increment on a black background
(Figure 2c), the main effect of interocular excursion
ratio was significant, F(2, 10) = 53.1, p < 0.001, ηp 2
= 0.914. In the same way as the conditions described
above, the base PSE was smaller for the equal (1:1) ratio
(11.7 cd/m2 ) than for the unequal ratios (15.7 and 14.8
cd/m2 for the 1:0.5 and 0.5:1 ratios, respectively). The
main effects of disparity and the interaction were not
significant (p > 0.05). Dunnett’s multiple-comparison
test showed no significant effect of binocular disparity
(p > 0.05).
Because the standard and comparison stimuli had
different shapes and appeared in a fixed temporal
order, the observers were always able to distinguish
 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 6

Observer 1 Observer 2

(a) Decrement on Gray BG

20 Zero Cross. Uncross.

15
PSE of the Comparison Stimulus Matched to the Standard Stimulus (cd/m²)

10
L L L R R R L L L R R R
5

0
1:1 1:0.5 0.5:1 1:1 1:0.5 0.5:1

(b) Increment on Gray BG

20

15

10
L L L R R R L L L R R R
5

0
1:1 1:0.5 0.5:1 1:1 1:0.5 0.5:1

(c) Increment on Black BG

20

15

10
L L L R R R L L L R R R
5

0
1:1 1:0.5 0.5:1 1:1 1:0.5 0.5:1

Interocular Excursion Ratio of the Comparison Stimulus

Figure 3. Brightness PSEs for two representative observers plotted separately in the left and right panels. The error bars indicate the
95% CIs of the estimated PSEs. (a) Results for the luminance decrement on a gray background. (b) Results for the luminance
increment on a gray background. (c) Results for the luminance increment on a black background.

those stimuli; thus, response bias could have caused standard deviation of PSE was only 1.21 cd/m2 for the
an overall shift of PSEs in individual data. However, decremental stimuli with zero disparity. The mean PSEs
this type of constant error is supposed to influence the at the equal ratio (1:1) were also comparable with the
measurements equally among different conditions and standard excursion, as described above. Taken together,
therefore is orthogonal to our experimental purpose. it is unlikely that response bias artificially created the
In addition, the six observers’ PSEs were distributed observed differences across conditions in the present
within a limited range of excursion; for example, the experiment.

Downloaded from jov.arvojournals.org on 03/23/2024

 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 7

Model fitting then divided by the denominator expression. Following

Kingdom and Libenson (2015) used a binocular
processing model to describe the relationship between
binocular color-saturation perception and dichoptic
stimuli with an identical hue but different saturation
values. We used their model to visualize the effects
of binocular disparity on binocular brightness
combination. The model is based on a model of
contrast normalization proposed by Legge and Foley
(1980) and its extensions for binocular processing
(Ding & Sperling, 2006; Maehara & Goryo, 2005;
Meese, Georgeson, & Baker, 2006). Although Ding
and Levi (2017) recently proposed a binocular
processing model that can explain various aspects of
binocular perception, including binocular brightness
combinations, its computations are much more complex
than the model used by Kingdom and Libenson (2015).
Therefore, for simplicity, we adopted their model as a
descriptive model for the present data, which can be
expressed by the following equation:
CLp CRp
B= +
z + CLq + wCRq z + CRq + wCLq
The right side of the equation consists of two terms
that represent two monocular responses. CL and CR
are the luminance excursions in the left and right eyes,
respectively. These monocular inputs are first raised
to the power of p for the numerator and q for the
denominator. The raised signals from the two eyes and
a constant (z) are summed in denominators. Before
this summation, the raised signal from the other eye is
multiplied by a weighting parameter (w) that controls
how much one eye is suppressed by the other eye. The
numerator expression in each monocular response is
the division, two monocular responses are summed to
yield binocular brightness (B).
The weighting parameter, w, is important for the
analysis of binocular luminance combinations. If the
weighting parameter is approximately 1, then the raised
signals from the two eyes are simply summed in the
denominator expressions. Because this process works
as interocular suppression, the eye receiving the higher
excursion predominates the binocular brightness when
the luminance is unbalanced between the two eyes. On
the other hand, if the weighting parameter is near 0,
then each monocular response is virtually free from
the divisive suppression from the other eye. Therefore,
binocular brightness is determined based on the linear
summation of the two monocular responses. See
Kingdom and Libenson (2015) for more details about
the weighting parameters.
We fit this descriptive model to the data using a
procedure similar to that of Kingdom and Libenson
(2015). Each PSE was normalized to the PSE at an equal
ratio (1:1). For example, if the luminance excursions in
the left and right eyes are 15 and 7.5 cd/m2 , respectively,
at the PSE for an unequal ratio (1:0.5) and if the PSE
for the equal ratio is 12 cd/m2 , then their normalized
values are 1.25 and 0.625, respectively. The normalized
luminance excursions in the two eyes, L and R, are at
the PSE when the binocular brightness (B) equals 1.
An equal brightness curve, which corresponds to the
family of PSEs to the standard stimulus predicted by
the model, can be plotted on the plane in which the x-
and y-axes indicate normalized luminance in the left
and right eyes, respectively. The model prediction was
constrained to pass through the PSE for an equal ratio
because its normalized value was one by definition
and a constant z was fixed to be near zero. Therefore,

wzero wcross wuncross RMSEzero RMSEcross RMSEuncross

Decrement on a Gray Background

Mean 0.780 0.631 0.675 0.0573 0.0531 0.0291
95% CI
Upper limit 0.985 0.689 0.785
Lower limit 0.575 0.573 0.565
Increment on a Gray Background
Mean 0.464 0.440 0.406 0.0550 0.0677 0.0251
95% CI
Upper limit 0.588 0.535 0.496
Lower limit 0.340 0.345 0.317
Increment on a Black Background
Mean 0.246 0.260 0.245 0.0814 0.0698 0.0654
95% CI
Upper limit 0.324 0.394 0.376
Lower limit 0.169 0.125 0.114
Table 1. Mean w Values, 95% CI Limits, and Mean RMSEs for Experiment 1.

Downloaded from jov.arvojournals.org on 03/23/2024

 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 8

each fit aimed to find the curve that ran closest to Table 1 presents the mean w values, their 95% CIs,
the two data points in the dichoptic conditions. The and mean RMSEs. The √ mean RMSEs were reasonably
model contained four parameters, but p, q, and z small compared with 2 of the radial distance to the
were fixed at 4.25, 3.0, and 0.001, respectively. These standard value, suggesting that the model fit the data
parameter values are the same as the constants used by well. The smooth curves in the left column of Figure 4
Kingdom and Libenson (2015). We used the MATLAB correspond to the model fits with interobserver mean
(MathWorks, Natick, MA) “fminbnd” function to find w values. In this figure, the normalized luminance was
the w value that minimized the root mean squared error reconverted back to cd/m2 before plotting. The best
(RMSE) between the PSE data and the predicted PSEs fit to the data from the two representative observers is
in the radial direction. The w-value range was set to be 0 shown in the middle and right columns (see Figure A1
to 1.25. in the Appendix for all individual data in the condition

Mean Parameters Observer 1 Observer 2

25 25 25
Decrement Decrement Decrement
20 20 20
Gray BG Gray BG Gray BG
15 15 15

10 10 10
Luminance Excursion in the Right Eye (cd/m²)

5 5 5
n=6
0 0 0
0 5 10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25

25 25 25
Increment Increment Increment
20 20 20
Gray BG Gray BG Gray BG
15 15 15

10 10 Zero 10

5 5 Cross. 5
n=6 Uncross.
0 0 0
0 5 10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25

25 25 25
Increment Increment Increment
20 20 20
Black BG Black BG Black BG
15 15 15

10 10 10

5 5 5
n=6
0 0 0
0 5 10 15 20 25 0 5 10 15 20 25 0 5 10 15 20 25

Luminance Excursion in the Left Eye (cd/m²)

Figure 4. Equal brightness curves for the decrement on a gray background (top panels), the increment on a gray background (middle
panels), and the increment on a black background (bottom panels). Smooth curves in the left panels correspond to the model curves
with mean w values for the zero-disparity (blue), crossed-disparity (red), and uncrossed-disparity (yellow) conditions. The blue dotted
lines represent the winner-take-all prediction (vertical and horizontal lines) and the linear-averaging prediction (diagonal line) in the
zero-disparity condition. The central and right panels show the fit to the individual data of two representative observers. Symbols in
the figure represent the PSEs. The error bars indicate the 95% CIs of the estimated PSEs. Most error bars are smaller than the symbols.

Downloaded from jov.arvojournals.org on 03/23/2024

 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 9

with the decrement on a gray background). The equal 30 30

brightness curves had greater curvatures, especially 25 Observer 5 25 Observer 6
at the center, for the luminance decrement on a gray 20 20
background (top row), compared to the conditions
15 15
of the luminance increments (middle and bottom Zero

Luminance Excursion in the Right Eye (cd/m²)

rows). The curves show a quasilinear function for 10 10
Cross.
the luminance increment on a black background 5
Uncross. 5
(bottom row). The curves for the luminance increment 0 0
on a gray background were in between (middle 0 5 10 15 20 25 30 0 5 10 15 20 25 30
row).
As noted previously, ANOVAs and Dunnett’s 30 30
multiple comparison tests of PSEs indicate that crossed 25 Observer 7 25 Observer 8
disparity had a significant effect on how the visual 20 20
system integrates luminance decrements presented to
15 15
the left and right eyes. This significant effect can be
observed as a change in the equal brightness curves 10 10
for the luminance decrement on a gray background 5 5
(top rows of Figure 4). The curves appear to be less 0 0
bent for the crossed disparity (red) than for the zero 0 5 10 15 20 25 30 0 5 10 15 20 25 30
disparity (blue). The mean w values reflect this trend
(Table 1a). This was smaller for the crossed disparity 30 30
(0.631) than for the zero disparity (0.780). Although 25 Observer 9 25 Observer 10
we subjected w values to Dunnett’s test, the difference 20 20
between zero and crossed disparity reached a statistical 15 15
significance level of p = 0.064, which is rather marginal.
10 10
However, this may be because three (p, q, and z) of
the four model parameters were fixed at the constants 5 5
used by Kingdom and Libenson (2015) and thus are 0 0
not guaranteed to be optimal for our observers. The 0 5 10 15 20 25 30 0 5 10 15 20 25 30
uncrossed disparity also yielded a smaller exponent Luminance Excursion in the Left Eye (cd/m²)
parameter (0.675) than zero disparity, resulting in
slightly less bent curves (Figure 4, yellow); however,
Figure 5. Equal brightness curves for luminance decrement on a
this trend was not significant in Dunnett’s test. For
gray background in Experiment 2. Each panel shows the fit to
the luminance increments on the gray and black
the individual data. Symbols represent the PSEs. Error bars
backgrounds (Figure 4, middle and bottom rows),
indicate the 95% CIs of the estimated PSEs. Most error bars are
the curves almost overlap. The mean w values were
smaller than the symbols.
comparable across the three binocular disparities under
these conditions (Tables 1b and 1c), indicating that the
binocular disparity had little effect when luminance
increments were presented. in which binocular combination at zero disparity is
apparently winner-take-all. According to Baker et
al. (2012) and Ding and Levi (2017), the binocular
combination of decrements is intermediate between
Results of Experiment 2: Luminance winner-take-all and linear averaging at small excursions
decrement with a larger excursion but gradually gets closer to winner-take-all with
increasing excursion. Therefore, in the next experiment,
In Experiment 1, the crossed disparity produced we used stimuli with a larger decremental excursion
a significant change in the binocular brightness (1.5 times) than those in Experiment 1. The levels of
combination of luminance decrement on a gray interocular excursion ratios were also increased from
background, although there was no significant effect of three to five for more precise estimations of equal
disparity for luminance increment on a gray and black brightness curves.
background. The effect of disparity may depend on Figure 5 illustrates the individual results for the
the pattern of binocular combination; that is, disparity six observers. As expected from the above-mentioned
had no effect on luminance increments because the literature, the equal brightness curves were more bent
binocular combination was already close to linear than those in Experiment 1, indicating that binocular
averaging at zero disparity. If this was the case, then the brightness was determined in a winner-take-all manner.
effect of disparity might become larger in a situation However, observer 10 showed a somewhat different

Downloaded from jov.arvojournals.org on 03/23/2024

 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami
wzero wcross
Mean 1.15 0.916
95% CI
Upper limit 1.26 1.18
Lower limit 1.04 0.656
Table 2. Mean w Values, 95% CI Limits, and Mean RMSEs for Decrement on a Gray Background in Experiment 2.
pattern (see the bottom-right panel in Figure 5). Even
when the excursion was larger for the left eye than for
the right eye, binocular combination suggested a strong
dominance of the right eye. Due to these asymmetrical
results between the two eyes, the model failed to fit the
data for observer 10. In addition, observer 10 exhibited
very large 95% CIs (error bars in Figure 5) compared
with those for the other observers. Notably, despite the
principle that the range should ideally be spanning 0%
to 100% to derive a reliable sigmoidal psychometric
function by best-fitting, inspection of the raw data
confirmed that in only five of the 15 conditions did the
response rates in the method of constant stimuli span
<25% to >75%. Due to these indications of atypical
binocular vision and poor brightness discriminability,
we excluded observer 10 from the following
analysis.
Comparisons of PSEs
We subjected the base PSE values to a two-way
repeated-measures ANOVA with the interocular
excursion ratio (1:1, 1:0.75, 1:0.5, 0.75:1, or 0.5:1) and
binocular disparity (zero, crossed, or uncrossed) as
factors. The base PSEs significantly differed across
interocular excursion ratios, F(4, 16) = 7.09, p < 0.001,
ηp 2 = 0.639. This significant main effect indicates
that the base PSE was smaller for the equal (1:1)
ratio (19.1 cd/m2 ) than for the dichoptic conditions
(20.4, 20.3, 20.2, and 20.2 cd/m2 for the 1:0.75, 1:0.5,
0.75:1, and 0.5:1 ratios, respectively). Although the
main effect of binocular disparity was not significant
(p > 0.05), the two factors significantly interacted,
F(8, 32) = 3.59, p < 0.001, ηp 2 = 0.473. To clarify
the interaction, we conducted Dunnett’s multiple
comparison test for each interocular ratio. The base
PSEs were significantly higher for the crossed disparity
than for zero disparity at the interocular excursion
ratios of 0.5:1 (20.7 cd/m2 vs. 19.6 cd/m2 ) and 0.75:1
(20.6 cd/m2 vs. 19.7 cd/m2 ). The 95% CIs of the
PSE elevations ranged from 0.410 to 1.75 cd/m2
for the 0.5:1 ratio (p = 0.005) and from 0.758 to
1.56 cd/m2 for the 0.75:1 ratio (p = 0.033). These
elevations in the PSEs are consistent with the results
of Experiment 1, although there was no significant
difference at the interocular excursion ratios of 1:0.5
and 1:0.75 (p > 0.05). The uncrossed disparity produced
Downloaded from jov.arvojournals.org on 03/23/2024
10
wuncross RMSEzero RMSEcross RMSEuncross
0.904 0.0427 0.0285 0.0213
1.11
0.700
no significant elevation in PSEs (p > 0.05), but the
base PSEs were significantly lower for the uncrossed
disparity (18.8 cd/m2 ) than for zero disparity (19.6
cd/m2 ) at the equal ratio (1:1). The 95% CIs of the
PSE reduction ranged from 0.152 to 1.37 cd/m2 (p =
0.018).
The crossed disparity caused some elevation of PSEs
in the dichoptic conditions but not in the equal-ratio
condition, resulting in the equal brightness curve
being less bent for the crossed disparity than for zero
disparity. In contrast, the uncrossed disparity caused
the reduction of PSEs at the equal ratio but not in
the dichoptic conditions. This result is also consistent
with the shape change in the equal brightness curve in
a manner similar to the above-described case of the
crossed disparity. The aforementioned shape changes
from winner-take-all to slightly less curvatures at
non-zero disparities can be seen in Figure 5, especially
in the curves of observers 5 and 9.
Model fitting
We fit the model to the PSE data to visualize the
changes in equal brightness curves using the same
procedure as that in Experiment 1. Table 2 presents
the mean w values, their 95% CIs, and mean RMSEs.
The mean RMSEs were comparable to those in
Experiment 1 and reasonably small. The smooth curves
in Figure 5 are the fits to the individual data. Figure 6
shows the model curves with the interobserver mean
w values. Consistent with the results of previous
studies (Baker et al., 2012; Ding & Levi, 2017), the
curvatures were greater in Experiment 2 (Figure 6) than
in Experiment 1 (Figure 4). Dunnett’s test confirmed
that the crossed and uncrossed disparities produced
smaller w values (0.916 and 0.904, respectively) than
those in the zero-disparity condition (1.15). The 95%
CIs of the difference ranged from 0.0841 to 0.381 for
the crossed disparity (p = 0.006) and from 0.0957 to
0.393 for the uncrossed disparity (p = 0.004). Because
of the reduction in w values, the curvatures were less
bent for the crossed and uncrossed disparities than for
zero disparity (Figures 5 and 6). These results indicate
that, in the situation of luminance decrement with
a larger excursion, binocular disparities weakened
interocular suppression, and consequently the binocular
combination was less of a winner-take-all scenario.
 Journal of Vision (2024) 24(2):4, 1–14
30
in the Right Eye (cd/m²)
Luminance Excursion
25
20
15
Zero
10
Cross.
5 increments on a gray background did not change. The
Uncross.
0 due to the reduction in interocular suppression because
0 5 10 15 20
Luminance Excursion
in the Left Eye (cd/m²)
Figure 6. The model fits with mean w values for the
zero-disparity (blue), crossed-disparity (red), and
uncrossed-disparity (yellow) conditions in Experiment 2. The
blue dotted lines represent the winner-take-all prediction
(vertical and horizontal lines) and the linear-averaging
prediction (diagonal line) in the zero-disparity condition.
Discussion
The present study examined how binocular disparity
changed binocular brightness combination patterns.
The results of this study were consistent with those of
previous studies in that luminance decrements in the
left and right eyes were combined in a winner-take-all
fashion, whereas the combination of luminance
increments was close to linear averaging when the
luminance values in the two eyes were not very different
(Anstis & Ho, 1998; Baker et al., 2012; Ding & Levi,
2017). Our main finding was that the binocular
brightness combination slightly deviated from the
winner-take-all prediction when a luminance decrement
was accompanied by a crossed disparity (Experiment 1)
or by a crossed or uncrossed disparity (Experiment 2).
In contrast, for luminance increments, binocular
disparity had no effect on the binocular brightness
combination.
It was expected that the presence of binocular
disparity would cause binocular brightness
combinations to approach linear averaging if stimuli
with non-zero binocular disparity weakened interocular
suppression. However, the effect of binocular disparity
was significant only for a luminance decrement and
not for increments. Previous studies have reported that
the monocular luminance values of decrements are
combined in a nonlinear fashion (that is, winner-take-
all), whereas those of increments are combined in a
linear fashion, except for the range corresponding to
Fechner’s paradox (Anstis & Ho, 1998; Baker et al.,
Downloaded from jov.arvojournals.org on 03/23/2024
Maehara, Wang, & Murakami 11
2012; Ding & Levi, 2017; Engel, 1970; Levelt, 1965). The
reduction in interocular suppression due to non-zero
n=5 disparities may be effective only when the binocular
brightness combination for stimuli with zero disparity
is strongly nonlinear in the first place. According to
Baker et al. (2012), the effect of luminance excursion is
asymmetrical between increments and decrements. In
their results, the equal brightness curves for decrements
became slightly shifted toward linear averaging as the
luminance excursion decreased, whereas the curves for
effects of excursion level on decrements could also be
25 30
interocular suppression is generally considered to
weaken as the luminance contrast of stimuli decreases
(Maehara, Huang, & Hess, 2009; Schor & Heckmann,
1989). That is, the results of Baker et al. (2012) are
arguably consistent with the present results in that
the reduction of interocular suppression is a viable
idea in both cases to explain the reduction of strong
nonlinearity in the binocular combination of luminance
decrements.
Moreover, the luminance contrasts of the stimuli
might have caused the difference between the results of
luminance decrements and increments. In Experiment 1,
we equated Weber contrasts between the decrement
and increment on a gray background. Consequently,
the Michelson contrast, (Lmax − Lmin )/(Lmax + Lmin ),
of the decremental standard (25%) was higher than that
of the incremental standard (17%). Because interocular
suppression is assumed to be greater for stimuli with
higher contrasts, the effects of binocular disparity
could increase for decrements more than for increments
due to the difference in the Michelson contrast.
Nonetheless, according to Ding and Levi (2017), who
examined binocular combinations within a wider range
of luminance excursions, incremental stimuli show little
change in the equal brightness curve among different
excursions when background luminance is sufficiently
high (16.2 cd/m2 ). In light of their finding, we expect
that the present results of the incremental conditions
would not change dramatically even if the luminance
contrast was increased. Future investigations with a
wider range of excursions with disparity manipulations
will resolve this interesting issue.
Lu and Sperling (2012) summarized psychophysical
studies concerning the asymmetry between luminance
increments and decrements by proposing that
black (decremental) stimuli are represented with
larger magnitudes in the visual system than white
(incremental) stimuli when they have the same
physical intensity in virtually all asymmetric cases.
The asymmetry between luminance increments and
decrements may reflect the characteristics of natural
images. Based on natural image statistics, researchers
have suggested that retinal images contain a greater
number of dark features than bright features (Cooper
 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami
& Norcia, 2015; Ratliff, Borghuis, Kao, Sterling, &
Balasubramanian, 2010). Notably, one study suggested
that non-zero binocular disparities appear more
frequently in dark features than in bright features in
natural images (Cooper & Norcia, 2015). Therefore, it is
possible that our visual system represents dark features
more dominantly than bright features during binocular
disparity processing. This asymmetry may have caused
a difference in the effects of binocular disparity between
luminance increments and decrements. Cooper and
Norcia (2015) reported that dark features were more
prominent at lower spatial frequencies. In our study, the
observers had to judge the brightness of the surfaces but
not the luminance contrast of the edges. Judgment of
the former depends on information from components
with relatively low spatial frequencies. This may be
another reason why the effect of binocular disparity
was significant only for the luminance decrement.
The effect of binocular disparity was significant
for crossed disparity but not for uncrossed disparity
in Experiment 1, although both types of disparity
produced significant effects in Experiment 2. Various
studies have demonstrated separate mechanisms that
selectively process crossed and uncrossed disparities
(see for a review, see Mustillo 1985). The results of
Experiment 1 presumably reflected the differential
characteristics of these mechanisms. Manning, Finlay,
Neill, and Frost (1987) found that the duration
threshold for detection was substantially longer for
stimuli with uncrossed disparity than for those with
crossed disparity. Patterson et al. (1995) also showed
that a longer duration was required to perceive depth
from an uncrossed disparity than from a crossed
disparity. These studies suggest that the mechanism
for crossed disparity is more sensitive than that
for uncrossed disparity. Although our stimuli were
suprathreshold and 100% detectable, they are only
briefly presented. The stimuli with crossed disparity
might have elicited a larger neural response than those
with uncrossed disparity in the present experiment,
causing a difference in the effectiveness of the binocular
brightness combination in Experiment 1.
In the present experiments, the largest excursion
difference between the two eyes was found in the
excursion ratios of 1:0.5 and 0.5:1. If we could measure
PSEs in even greater unbalanced ratios, we could
make a more precise evaluation of the nonlinearity
of equal-brightness data. We tested the interocular
excursion ratios of L:R = 1:0.25 and 0.25:1 in a
preliminary experiment. When non-zero disparities
were added, however, observers were not able to fuse
the stimuli as in cyclopean vision but instead saw
only two dichoptic images, one brighter and the other
darker, making it difficult to perform the brightness
task as in the main experiment. Therefore, our PSE
measurements had to be limited to range between 1:0.5
and 0.5:1 cases. One possible extension of the stimuli is
Downloaded from jov.arvojournals.org on 03/23/2024
12
to add a high-contrast rectangular contour to support
binocular fusion of our comparison stimulus, whereas
the luminance inside the contour varies for each eye.
This will enable us to examine the effects of binocular
disparity under conditions of greater luminance
differences between the eyes. However, in such cases,
the luminance of the contour can affect the brightness
perception of the interior. In addition, a process of
perceptual filling-in may be triggered by the spatial
profiles of luminance around the contour. Therefore,
elucidating the effects of contours on the binocular
brightness combination requires future studies.
Maehara and Murakami (2020) reported that
perceived contrast was approximately 1.1 times higher
at 1° binocular disparity than at zero disparity. If this
perceptual enhancement had occurred, the position
of the equal brightness curves would shrink in the
lower-left direction when the stimuli had a non-zero
binocular disparity. However, this effect was not
observed in Experiment 1 (Figure 4) and was smaller in
Experiment 2 (Figure 6) than that reported by Maehara
and Murakami (2020). The absence or reduction of
perceptual enhancement may be owing to differences in
the range of binocular disparities. We presented a short,
thin bar as a stimulus for the binocular brightness
combination and set its binocular disparity within the
fusion limit. This is because brightness judgment in
double vision is difficult under dichoptic conditions.
In contrast, the Gabor patches used in the study by
Maehara and Murakami (2020) presented disparities
over the fusion limit, but within the stereopsis limit.
Another possible explanation for the above discrepancy
is that binocular luster is involved in the perceptual
enhancement. Previous studies have suggested that
its occurrence depends on stimulus size (for a review,
see Wendt & Faul, 2022). The size of the comparison
stimulus in the present study was arguably smaller
than the lower limit of binocular luster, whereas the
Gabor patches used in the study by Maehara and
Murakami (2020) were sufficiently large. Nonetheless,
the perceptual enhancement observed in their study
was also vigorous when the left-eye and right-eye
images were vertically disparate with virtually no
spatial overlap, which is an unfavorable situation for
luster.
Conclusions
We found that, although the binocular brightness
combination of the luminance decrement tended
to be nonlinear and more like winner-take-all, the
combination became less of a winner-take-all scenario
when presented with non-zero disparity. This shift
in binocular combination is possibly due to weaker
interocular suppression caused by non-zero binocular
 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami
disparity. This is presumably because simultaneous
stimulations at corresponding retinal points are required
for strong suppression. Another possible mechanism
is that stereopsis, which is processed when disparity is
signaled, inhibits interocular suppression to avoid eye
rivalry and to promote binocular fusion, resulting in the
maximal use of precise position information originating
from two monocular signals. The mechanism of
attenuation of the interocular suppression we found still
remains unclear, and we argue that one of the possible
extensions would be to introduce vertical disparity to
ascertain whether the attenuation requires the process
for stereopsis specifically or just any misalignment from
the corresponding retinal points.
Previous studies have suggested that interocular
suppression is reduced in Ganzfeld stimuli (Bourassa
& Rule, 1994) and balanced between the two eyes in
binocularly matched features (Jennings & Kingdom,
2019; Kingdom & Libenson, 2015; Kingdom &
Wang, 2015; Wang et al., 2022). These techniques
could be effective tools for training binocular vision
in stereoblind observers or patients with amblyopia.
Here, we suggest that binocular disparity manipulation
potentially offers another technique to make both eyes
functional. However, the effects of binocular disparity
were small and limited to a luminance decrement in
the present experiment. For such training purposes,
future studies are required to determine the optimal
stimulus conditions to elicit the largest attenuation
effect on interocular suppression in the domain of
brightness.
Keywords: brightness, stereopsis, interocular
suppression, binocular perception
Acknowledgments
IM was supported by a JSPS KAKENHI
Grant-in-Aid for Scientific Research (21H04426).
Commercial relationships: none.
Corresponding author: Goro Maehara.
Email: goro@kanagawa-u.ac.jp.
Address: Department of Human Science, Kanagawa
University, Kanagawa, Japan.
* binocular luminance contrast. Vision Research, 116,
GM and YW contributed equally to this work.
References
Anstis, S., & Ho, A. (1998). Nonlinear combination of
luminance excursions during flicker, simultaneous
contrast, afterimages and binocular fusion. Vision
Research, 38(4), 523–539.
Downloaded from jov.arvojournals.org on 03/23/2024
13
Baker, D. H., Wallis, S. A., Georgeson, M. A., &
Meese, T. S. (2012). Nonlinearities in the binocular
combination of luminance and contrast. Vision
Research, 56, 1–9.
Blake, R. (1989). A neural theory of binocular rivalry.
Psychological Review, 96(1), 145.
Blake, R., & Fox, R. (1973). The psychophysical
inquiry into binocular summation. Perception &
Psychophysics, 14(1), 161–185.
Bourassa, C. M., & Rule, S. J. (1994). Binocular
brightness: A suppression-summation trade off.
Canadian Journal of Experimental Psychology,
48(3), 418–434.
Cooper, E. A., & Norcia, A. M. (2015). Predicting
cortical dark/bright asymmetries from natural
image statistics and early visual transforms. PLoS
Computational Biology, 11(5), e1004268.
Ding, J., & Levi, D. M. (2017). Binocular combination
of luminance profiles. Journal of Vision, 17(13):4,
1–32, https://doi.org/10.1167/17.13.4.
Ding, J., & Levi, D. M. (2021). A unified model for
binocular fusion and depth perception. Vision
Research, 180, 11–36.
Ding, J., & Sperling, G. (2006). A gain-control theory of
binocular combination. Proceedings of the National
Academy of Sciences, USA, 103(4), 1141–1146.
Engel, G. (1970). Tests of a model of binocular
brightness. Canadian Journal of Psychology, 24(5),
335–352.
Fechner, G. T. (1860). Elemente der Psychophysik.
Leipzig, Germany: Breitkopf & Härtel.
Jennings, B. J., & Kingdom, F. A. A. (2019). Unmasking
the dichoptic mask. Journal of Vision, 19(6):3, 1–8,
https://doi.org/10.1167/19.6.3.
Kingdom, F. A. A., & Libenson, L. (2015). Dichoptic
color saturation mixture: Binocular luminance
contrast promotes perceptual averaging. Journal of
Vision, 15(5):2, 1–15, https://doi.org/10.1167/15.5.2.
Kingdom, F. A. A., & Prins, N. (2016). Psychophysics:
A practical introduction (2nd ed.). Cambridge, MA:
Academic Press.
Kingdom, F. A. A., & Wang, D. (2015). Dichoptic
colour-saturation masking is unmasked by
45–52.
Kingdom, F. A. A., Yared, K.-C., Hibbard, P. B., &
May, K. A. (2020). Stereoscopic depth adaptation
from binocularly correlated versus anti-correlated
noise: Test of an efficient coding theory of
stereopsis. Vision Research, 166, 60–71.
Legge, G. E., & Foley, J. M. (1980). Contrast masking
in human vision. Journal of the Optical Society of
America, 70(12), 1458–1471.
 Journal of Vision (2024) 24(2):4, 1–14 Maehara, Wang, & Murakami 14

Lema, S. A., & Blake, R. (1977). Binocular summation on stereopsis and fusion. Vision Research, 29(7),
in normal and stereoblind humans. Vision Research, 837–847.
17(6), 691–695. Wang, M., Ding, J., Levi, D. M., & Cooper, E. A.
Levelt, W. J. (1965). On binocular rivalry (dissertation, (2022). The effect of spatial structure on binocular
Leiden University). Berlin: De Gruyter Mouton. contrast perception. Journal of Vision, 22(12):7,
Lu, Z.-L., & Sperling, G. (2012). Black–white 1–19, https://doi.org/10.1167/jov.22.12.7.
asymmetry in visual perception. Journal of Vision, Wendt, G., & Faul, F. (2022). Binocular luster – A
12(10):8, 1–21, https://doi.org/10.1167/12.10.8. review. Vision Research, 194, 108008.
Maehara, G., & Goryo, K. (2005). Binocular,
monocular and dichoptic pattern masking. Optical
Review, 12(2), 76–82. Appendix
Maehara, G., Huang, P.-C., & Hess, R. F. (2009).
Importance of phase alignment for interocular
suppression. Vision Research, 49(14), 1838–1847. 25 25
Maehara, G., & Murakami, I. (2020). Perceptual Observer 1 Observer 2
20 20
enhancement of suprathreshold luminance
modulation in stereoscopic patterns. Journal of 15 15
Vision, 20(12):8, 1–11, https://doi.org/10.1167/jov. 10 10
20.12.8. Zero

Luminance Excursion in the Right Eye (cd/m²)

Manning, M. L., Finlay, D. C., Neill, R. A., & Frost, 5 Cross. 5
B. G. (1987). Detection threshold differences to 0
Uncross. 0
crossed and uncrossed disparities. Vision Research, 0 5 10 15 20 25 0 5 10 15 20 25
27(9), 1683–1686.
May, K., & Zhaoping, L. (2022). Li and Atick’s 25 25
theory of efficient binocular coding: A tutorial and 20
Observer 3
20
Observer 4
mini-review. Vision Research, 201, 107950.
Meese, T. S., Georgeson, M. A., & Baker, D. H. 15 15
(2006). Binocular contrast vision at and above 10 10
threshold. Journal of Vision, 6(11):7, 1224–1243,
https://doi.org/10.1167/6.11.7. 5 5

Mustillo, P. (1985). Binocular mechanisms mediating 0 0

crossed and uncrossed stereopsis. Psychological 0 5 10 15 20 25 0 5 10 15 20 25
Bulletin, 97(2), 187–201.
Patterson, R., Cayko, R., Short, G. L., Flanagan, R., 25 25
Moe, L., Taylor, E., . . . Day, P. (1995). Temporal 20
Observer 5
20
Observer 6
integration differences between crossed and
uncrossed stereoscopic mechanisms. Perception & 15 15
Psychophysics, 57, 891–897. 10 10
Ratliff, C. P., Borghuis, B. G., Kao, Y.-H., Sterling, P.,
5 5
& Balasubramanian, V. (2010). Retina is structured
to process an excess of darkness in natural scenes. 0 0
Proceedings of the National Academy of Sciences, 0 5 10 15 20 25 0 5 10 15 20 25
USA, 107(40), 17368–17373.
Luminance Excursion in the Left Eye (cd/m²)
Rose, D., Blake, R., & Halpern, D. L. (1988). Disparity
range for binocular summation. Investigative
Figure A1. Individual results for the decrement on a gray
Ophthalmology & Visual Science, 29(2), 283–290.
background in Experiment 1. Smooth curves in the left panels
Said, C. P., & Heeger, D. J. (2013). A model of binocular correspond to the model fit with mean w values for the
rivalry and cross-orientation suppression. PLoS zero-disparity (blue), crossed-disparity (red), and
Computational Biology, 9(3), e1002991. uncrossed-disparity (yellow) conditions. Symbols in the figure
Schor, C., & Heckmann, T. (1989). Interocular represent the PSEs. Error bars indicate the 95% CIs of the
differences in contrast and spatial frequency: Effects estimated PSEs. Most error bars are smaller than symbols.

Downloaded from jov.arvojournals.org on 03/23/2024