Marta Suárez-Pinilla1, Anil K Seth1, Warrick Roseboom1. 1. Sackler Centre for Consciousness Science, University of Sussex, Brighton, UK; Department of Informatics, University of Sussex, Brighton, UK.
Abstract
Visual experience appears richly detailed despite the poor resolution of the majority of the visual field, thanks to foveal-peripheral integration. The recently described uniformity illusion (UI), wherein peripheral elements of a pattern take on the appearance of foveal elements, may shed light on this integration. We examined the basis of UI by generating adaptation to a pattern of Gabors suitable for producing UI on orientation. After removing the pattern, participants reported the tilt of a single peripheral Gabor. The tilt aftereffect followed the physical adapting orientation rather than the global orientation perceived under UI, even when the illusion had been reported for a long time. Conversely, a control experiment replacing illusory uniformity with a physically uniform Gabor pattern for the same durations did produce an aftereffect to the global orientation. Results indicate that UI is not associated with changes in sensory encoding at V1 but likely depends on higher level processes.
Visual experience appears richly detailed despite the poor resolution of the majority of the visual field, thanks to foveal-peripheral integration. The recently described uniformity illusion (UI), wherein peripheral elements of a pattern take on the appearance of foveal elements, may shed light on this integration. We examined the basis of UI by generating adaptation to a pattern of Gabors suitable for producing UI on orientation. After removing the pattern, participants reported the tilt of a single peripheral Gabor. The tilt aftereffect followed the physical adapting orientation rather than the global orientation perceived under UI, even when the illusion had been reported for a long time. Conversely, a control experiment replacing illusory uniformity with a physically uniform Gabor pattern for the same durations did produce an aftereffect to the global orientation. Results indicate that UI is not associated with changes in sensory encoding at V1 but likely depends on higher level processes.
Visual experience appears richly detailed despite the poor sensory precision of the
majority (periphery) of the visual field. This topic has received considerable
recent attention (Cohen, Dennett,
& Kanwisher, 2016; Haun, Tononi, Koch, & Tsuchiya, 2017), with debate about the degree
to which visual experience is in fact rich and the potential perceptual processes
that may contribute to apparent richness. One recent study demonstrated a compelling
example of how the rich detail within the high-precision central visual field alters
peripheral perception—the uniformity illusion (UI; Otten, Pinto, Paffen, Seth, & Kanai,
2016). UI describes a phenomenon wherein apparent perceptual uniformity
occurs when variable sensory stimulation is presented in peripheral vision, while
the central visual field is presented with uniform stimuli. UI occurs for a wide
variety of perceptual dimensions, including relatively low-level sensory features
such as orientation or colour and higher level features such as density (see
www.uniformillusion.com for examples).We sought to examine the mechanisms underlying UI using perceptual adaptation. It is
well established that exposure to a specific stimulus magnitude (like an oriented
grating) causes perceptual aftereffects (e.g., tilt aftereffect [TAE]; Gibson & Radner, 1937).
For visual orientation, perceptual aftereffects have been associated with specific
changes in neural coding at the primary visual cortex (V1) and are localized in a
retinotopic reference frame (Knapen, Rolfs, Wexler, & Cavanagh, 2010). Here, we utilize the
spatial specificity of TAE to examine whether the apparent perceptual uniformity in
UI can be attributed to changes in V1-based neural coding for visual orientation.
Specifically, we presented participants with Gabor grids wherein the orientation of
central elements was uniform, but the orientation of peripheral elements was
variable—producing UI. At fixed test locations in the periphery of the grid, we
presented a physical orientation that differed from the global illusory percept,
thus putting local and global orientation in opposition. Following prolonged
exposure to global illusory uniformity (UI), we contrasted whether the resultant TAE
was consistent with the local, physical orientation or the illusory global
orientation.
Methods
Procedure
The experiment had two parts: illusion session and control session. Each session
contained six blocks, and each block had an adaptation phase and a test phase
(Figure 1). A
practice block was run before the illusion session to familiarize participants
with UI.
Figure 1.
Experimental structure. During the adaptation phase, participants
were presented with a Gabor grid wherein the central Gabors had a
uniform orientation, while peripheral orientations were
heterogeneous. Under UI, perceptual experience was that of a uniform
pattern with all Gabors tilted like the central ones. This illusory
percept alternated with a nonillusory, nonuniform percept at
different times during adaptation. For a specific peripheral Gabor
(adapting Gabor), physical and illusory orientation were always in
opposition. The control session replicated the phenomenology of the
illusion session, replacing perceived with physical uniformity at
times in which the participant reported UI in the illusion session.
The test phase had 24 trials, wherein participants reported the tilt
of a single peripheral Gabor whose location coincided with the
adapting Gabor.
Experimental structure. During the adaptation phase, participants
were presented with a Gabor grid wherein the central Gabors had a
uniform orientation, while peripheral orientations were
heterogeneous. Under UI, perceptual experience was that of a uniform
pattern with all Gabors tilted like the central ones. This illusory
percept alternated with a nonillusory, nonuniform percept at
different times during adaptation. For a specific peripheral Gabor
(adapting Gabor), physical and illusory orientation were always in
opposition. The control session replicated the phenomenology of the
illusion session, replacing perceived with physical uniformity at
times in which the participant reported UI in the illusion session.
The test phase had 24 trials, wherein participants reported the tilt
of a single peripheral Gabor whose location coincided with the
adapting Gabor.
Illusion session
Each block began with an adaptation phase, in which participants were
presented with a grid of Gabor patches suitable for producing the UI,
affecting the apparent orientation of peripheral elements: All Gabors in the
central area had a uniform orientation, whereas orientation of the
peripheral Gabors was heterogeneous. Gaze-contingent stimulus presentation
ensured that each Gabor was presented to a specific retinal location, as the
entire pattern was removed if the participant’s gaze deviated from central
fixation by more than 1.5 degrees of visual angle (dva)—a tolerance
threshold equivalent to half the size of each cell of the grid. Adaptation
lasted 180 seconds, but because the stimulus was removed when fixation
lapsed, actual exposure time could be shorter.Participants reported the experience of illusory uniformity by pressing a key
when all Gabors appeared to take a uniform orientation.The test phase had 24 trials, separated by a pseudorandom interval of 1,000
to 1,500 ms. In each trial, a single Gabor (test Gabor) was presented for
500 ms at a specific peripheral location, coinciding with the position of a
specific Gabor during adaptation (adapting Gabor). Participants reported if
the test Gabor was tilted clockwise (CW) or counterclockwise (XCW) from
vertical.
Control session
The control session also had six blocks, each built to replicate the
phenomenology of a homologous block of the illusion session but replacing
illusory for physical uniformity during the adaptation phase.During the adaptation phase in the illusion session, an empty background was
presented whenever the gaze-contingent mechanism removed the adapting
pattern. The same pattern of stimulus presentation and removal was
replicated in the control session. The stimulus was additionally removed
whenever fixation lapsed in the control session. At any other time, the
presentation displayed one of two patterns, differing only in the
orientation of peripheral Gabors. The first was identical to the pattern
presented in the illusion session and was displayed at times in which the
participant had not reported UI during adaptation in the
illusion session. At times during which the participant had reported UI, the
presented pattern was one in which all Gabors had the same
physical orientation, consistent with the desired
illusory orientation during the illusion session. Thus, physical uniformity
was inserted at the times in which illusory uniformity had been reported in
the illusion session. Participants were not informed that this would
occur.The test phase was identical to that in the illusion session: The location
and orientation of the test Gabor in each trial was identical, as well as
its test latency (time between the end of the adaptation phase and stimulus
onset).
Stimuli
Stimuli were displayed on dark grey background (1.96 cd/m2). A red
fixation dot (8.34 cd/m2, 0.42 dva diameter) showed constantly on the
screen centre.
Gabor patches
Each Gabor consisted of a sine-wave luminance grating with Michelson contrast
of 1, 0° phase, and spatial frequency of 1.66 cycles per dva, and a two
dimensional Gaussian envelope with a sigma of 0.43 dva.
Adapting pattern
The adapting pattern spanned the entire screen and consisted of a 13 × 17
grid formed by invisible square cells measuring 3 dva per side (Figure 1). Each Gabor
was presented in the centre of each cell. The central area spanned 15 dva
horizontally and vertically, encompassing all cells belonging to rows 5 to 9
and columns 7 to 11. All central Gabors had the same orientation, which
could be one of two values, each for half the blocks of one session: −15°
(global clockwise tilt [GCW]) or 15° (global counterclockwise tilt [GXCW]).
The orientations of peripheral Gabors were sampled from a discrete uniform
distribution centred on the global orientation and ranging 70° (35° to each
side). Thus, mean orientation was the same for central and peripheral Gabors
and matched the global orientation perceived under UI.Two peripheral Gabors of the pattern (adapting Gabors) corresponded to the
positions in which the test Gabors would be displayed during the test phase:
They were located along the middle (7th) row, at 12.02 dva left and right of
the screen centre (columns 5 and 13). Both had the same nonrandomized local
orientation, which was the opposite of the global orientation of the block:
either 15° (local counterclockwise tilt [LXCW]) or −15° (local clockwise
[LCW]).Henceforth, we give the label adapting condition CX to the presentation
pattern wherein the local orientation of the adapting Gabor is clockwise,
and the global orientation of the pattern is counterclockwise (LCW, GXCW).
Conversely, we will refer to the pattern with LXCW and GCW orientations as
adapting condition XC. Both conditions occurred equally frequently during
the experiment.As described above, during the control session, the adapting pattern was
replaced by a physically uniform pattern at those times during which
participants had reported UI in the illusion session. In these instances,
every Gabor in the pattern (including the adapting
Gabors) took the global orientation.
Test Gabors
A single test Gabor was presented per trial, matching the position of one of
the two adapting Gabors. Test Gabors were displayed in the left and right
hemifield with equal frequency per block and could take one of eight equally
frequent orientations: −12°, −5°, −2°, −1°, 1°, 2°, 5°, and 12° (negative
values indicate CW tilt). Thus, test orientations were always intermediate
between global and local orientations (−15°, 15°).
Participants
Participants were recruited through online advertisement, were older than 18
years, and reported normal or corrected-to-normal vision. This study received
ethical approval by the Research Ethics Committee of the University of
Sussex.
Apparatus
Experiments were programmed in MATLAB 2016a (MathWorks Inc., Natick, MA, USA) and
displayed on a LaCie Electron 22BLUE II 22″ with screen resolution of
1,024 × 768 pixels and refresh rate of 100 Hz. Eye-tracking was performed with
EyeLink 1000 Plus (SR Research, Mississauga, Ontario, Canada) at sampling rate
of 1000 Hz, with level desktop camera mount. Head position was stabilized 43 cm
from the screen using chin and forehead rest. Calibration of the eye-tracker was
performed at the beginning of each block with a standard five-point grid and a
maximal average error of 0.5 dva.
Statistical Analysis
Psychometric curve fitting was performed in MATLAB 2017b, using Palamedes
toolbox, version 1.8.1 (Prins & Kingdom, 2009). A cumulative Gaussian curve was fitted
by the method of maximum likelihood to the proportion of “XCW” responses per
test Gabor orientation, separately for each participant and session/condition
(depending on the specific analysis). The threshold (α) for 0.5 proportion of
XCW responses and the slope of the curve (β) were free parameters (starting
values: α = 0°, β = .04), while guessing (γ) and lapse rate (λ) were fixed at
zero.Bayesian statistics were conducted on JASP (JASP Team, 2017, version 0.8.3.1).
For Bayesian t tests, we employed as prior distribution
Cauchy(0, ) for two-sided predictions, or a folded Cauchy(0,
) for one-sided predictions (Measure 1 < Measure 2).
Likewise, for Bayesian Pearson correlations, we employed a uniform distribution
U(−1,1) for two-sided or U(0,1) for one-sided (positive) predictions. For each
contrast result, the prior utilized is indicated by the formulated prediction
and the subscripts in BF10 (two-sided) or BF−0
/BF+0 (one-sided).
Results
Thirty participants volunteered for the experiment: 23 female; mean age was 21.6
years. To ensure sufficient exposure to the adapting pattern, we excluded blocks
wherein the pattern had been displayed for less than two thirds of the adaptation
phase (<120 seconds), due to gaze-contingent stimulus removal. In such cases, the
corresponding blocks from both control and illusion sessions were removed to
maintain balance. This caused exclusion of 32.78% blocks (118/360), including the
entire data sets from five participants. Furthermore, because our analyses compared
responses across adapting conditions (CX/XC), two additional participants were
excluded as all their valid blocks were of only one condition. Results presented
here correspond to the remaining 23 participants. Overall, results for all 25
participants with valid blocks were very similar to this counterbalanced sample (see
Supplementary Materials, Section S5).
Adaptation Phase
Average exposure time to the adapting pattern per block was 164.13 and 149.47
seconds for the illusion and control sessions: 91.18% and 83.04% of the
adaptation phase, respectively. The lower proportion in the control session was
expected as pattern removal occurred whenever it had in the illusion session, in
addition to times of improper fixation in the control block.Perceived uniformity was reported, on average, for 43.48 seconds in the illusion
session, 26.77% of the time of pattern presentation (minimum 0.55%, maximum
72.23%). The proportion of time of perceived uniformity during the control
session was similar to that for the illusion session: 28.41% (minimum 0.59%,
maximum 78.42%, Bayesian paired-samples t test:
BF01 = 2.733—anecdotal evidence for the null hypothesis). Physical
uniformity in the control session was reported as perceptually uniform 68.13% of
the time; by contrast, the nonuniform pattern was reported as uniform only 9.24%
of the time. Possibly, presentation of a truly uniform pattern at times shifted
a subjective criterion for uniformity by comparison, leading to more
conservative reports in the control sessions.
Hypotheses and Measurements
The experiment placed adaptation to illusory and physical orientation in
opposition to disambiguate between two competing hypotheses: To decide between hypotheses, data were analysed to ascertain the
direction of the adaptation-induced bias. We calculated the proportion of XCW
reports per test Gabor orientation and obtained the best fitting cumulative
Gaussian psychometric curve. The point of subjective equality (PSE) was defined
as the test orientation at which 50% reports are XCW. Because CW orientations
have (conventionally) negative sign and vice versa, negative PSE indicates a XCW
bias, and positive PSE indicates a CW bias.The perceived orientation under UI has no effect on tilt adaptation;
the TAE is driven solely by the physical orientation of the adapting
Gabor.The global orientation perceived for the entire pattern (including
the adapting Gabor) under UI can produce a TAE.During the illusion session, a TAE driven by (i.e., away from) the local
orientation of the adapting Gabor would imply physical adaptation, while a
global-driven TAE would indicate adaptation to illusory orientation. During the
control session, both local- and global-driven TAE are compatible with physical
adaptation, as the adapting Gabor physically takes the global orientation at
times of reported illusory uniformity in the illusion session.By calculating participants’ PSE per adapting condition, we obtained two
measurements:PSECX and PSEXC. For a local-driven TAE,
responses for adapting condition CX should exhibit a XCW bias
compared with condition XC (PSECX < PSEXC),
and the reverse should happen for a global-driven TAE.dPSE = PSECX–PSEXC. We employ this as a summary
measure indicating the overall direction of the bias. A negative
dPSE indicates a predominance of local-driven TAE
(PSECX < PSEXC) consistent with
physical adaptation to the local orientation, while a positive dPSE
indicates a global-driven TAE, consistent with adaptation to the
illusion (or to the physical replication of the illusion during the
control session).
TAE Is Driven by Physical, Not Illusory Orientation
Overall effect
Illusion session
Figure 2(a)
presents the average proportion of XCW reports per test Gabor
orientation during the illusion session, separated by adapting condition
(CX or XC). For illustration purposes, it shows cumulative Gaussian
curves fitted to the pooled data. However, for analysis, we fitted each
participant’s responses separately: Individual fits are detailed in the
Supplementary Materials, Section S2. Individual PSEs for each adapting
condition are summarized in Figure 2(b). On average,
PSECX = −0.502° and PSEXC = 0.687° reflected a
XCW and CW bias, respectively: dPSE = −1.197°
(PSECX < PSEXC Bayesian paired-samples
t test: BF−0 = 3.057) indicated a local,
physical-driven adaptation.
Figure 2.
Response patterns by adapting condition. (a) and (c) presents
the sample’s proportion of counterclockwise (XCW) reports
per test Gabor orientation, separated by adapting condition,
during the illusion (a) and control (c) sessions. The dotted
lines show the best cumulative Gaussian fit for the
psychometric curve of each condition, fitted on the sample’s
pooled data (N = 23). (a) and (c) are
included for illustrative purposes only, as the PSEs
obtained for analysis were computed separately for each
participant’s data: These results are depicted in (b) and
(d), where the bar heights represent the average point of
subjective equality (PSE), computed separately per
participant, session, and condition. The error bars
represent the between-participant standard error. (b)
Illusion session. PSEs for both adapting conditions reflect
a bias away from local orientation
(local-driven TAE). (d) Control session. On average (d),
responses show a global-driven TAE in CX and are unbiased in
XC. These results show that perceived (illusion) and
physical (control) uniformity behave differently, suggesting
that the TAE is always driven by the physical orientation,
even when that orientation is unseen under UI.
Response patterns by adapting condition. (a) and (c) presents
the sample’s proportion of counterclockwise (XCW) reports
per test Gabor orientation, separated by adapting condition,
during the illusion (a) and control (c) sessions. The dotted
lines show the best cumulative Gaussian fit for the
psychometric curve of each condition, fitted on the sample’s
pooled data (N = 23). (a) and (c) are
included for illustrative purposes only, as the PSEs
obtained for analysis were computed separately for each
participant’s data: These results are depicted in (b) and
(d), where the bar heights represent the average point of
subjective equality (PSE), computed separately per
participant, session, and condition. The error bars
represent the between-participant standard error. (b)
Illusion session. PSEs for both adapting conditions reflect
a bias away from local orientation
(local-driven TAE). (d) Control session. On average (d),
responses show a global-driven TAE in CX and are unbiased in
XC. These results show that perceived (illusion) and
physical (control) uniformity behave differently, suggesting
that the TAE is always driven by the physical orientation,
even when that orientation is unseen under UI.
Control session
In the control session, the adapting Gabor physically takes the global
orientation of the pattern during times of reported uniformity in the
illusion session. If adaptation is produced by physical orientation, we
should observe a more global-driven TAE compared with the illusion
session: dPSEIL < dPSECO. Conversely, if
perceived orientation under UI causes adaptation, we should not see a
difference between perceived and physical uniformity:
dPSEIL = dPSECO.Results indicate predominance of global-driven TAE during the control
session (Figure
2(d)): PSECX-CO = 0.433°,
PSEXC-CO = −0.083°, dPSECO = 0.516°. A Bayesian
paired-samples t test comparing dPSE in both sessions
was consistent with physical-driven adaptation:
dPSEIL < dPSECO, BF−0 = 7.476.
Therefore, the absence of a global-driven TAE in the illusion session
was not simply because the global pattern of orientation was
insufficient to induce TAE; rather, the illusory (but not the physical)
global pattern was insufficient to induce TAE.The overall predominance of global-driven TAE in the control session,
despite presentation of the uniform pattern for only ∼27% of time, may
be related to a putatively stronger adaptation during this time due to
the adjacent Gabors, which then take the global orientation,
contributing to the receptive field(s) where the test Gabor will be
later presented. Note, however, that the size of each grid cell (3 dva)
is larger than the diameter of most receptive fields at V1 (around 1
dva; Bentley &
Salinas, 2009), and the relationship between stimulus size
and TAE strength is not always intuitive (Harris & Calvert, 1985;
Parker,
1972). Another possibility involves extraclassical receptive
field effects exerted by the global surround on the adapting Gabor when
the latter takes the global orientation (iso-orientation surround
suppression; Chen,
Chen, Gao, Yang, & Yan, 2015). Whatever the contribution
of these effects, they act differently on physical compared with
illusory iso-orientation, in the manner expected for low-level
processing of the former, but not the latter.In the Supplementary Materials (Sections S2 and S4), we reanalyse the
data set based on raw responses, rather than PSE from fitted curves.
Both approaches show the same pattern of results, indicating that
choices related to curve fitting and goodness-of-fit of psychometric
curves do not significantly affect our analyses.
Time-dependent effect
Overall, responses in the illusion and control session fit the hypothesis
that TAE under UI is only driven by physical and not
illusory orientation. However, in the illusion session, UI is perceived
during only ∼27% of pattern exposure, on average. Thus, it could be argued
that a global, illusion-driven TAE might have been present, but undetected
in the overall results, overshadowed by the local-driven TAE at times when
UI is not perceived. This possibility seems unlikely because responses in
the control session (with uniformity also presented ∼27% of time)
do show an influence of the global-driven TAE. Thus,
such a possibility could only hold if the TAE driven by illusory orientation
was weaker than that caused by physical orientation. To rule out this
possibility, we examined the data from the illusion sessions for evidence of
exposure time-dependency of the TAE magnitude. Because the TAE is
time-dependent (Patterson, Wissig, & Kohn, 2013), if illusion-driven
adaptation was in fact present, we should find evidence for a shift towards
more global/less local TAE with longer times of perceived uniformity.If the TAE is driven only by physical orientation, in
the illusion session, we should expect independence from time of
perceived uniformity. Conversely, if the perceived orientation under UI
causes adaptation, the response pattern should shift from predominantly
local-driven towards more global-driven for longer time of perceived
uniformity. We can assess this potential shift by examining dPSE. As
stated above, negative dPSE indicates predominance of local-driven TAE
and positive dPSE global-driven TAE. Thus, in the presence of
illusion-driven adaptation, dPSE should correlate positively with time
of perceived uniformity.As time measure, we employed the proportion of perceived uniformity (over
time of pattern presentation) for conveying the balance between local
and (putative) global effects. We analysed the bivariate correlation
between time of perceived uniformity and dPSE (Figure 3(c)). Pearson’s
correlation coefficient and 95% credible intervals were
r = −.199 (−.537, .219), with moderate evidence
against a positive correlation: BF+0 = 0.146.
Figure 3.
TAE by time of uniformity. Physical, but not
perceived uniformity, causes a shift towards global-driven
TAE in a time-dependent manner. (a and b) Participants are
classified into two groups according to whether their
average time of uniformity is below (lower half) or above
(upper half) the sample’s median and depict each group’s
average responses by adapting condition in the illusion (a)
and control (b) session. For illustration purposes, a
psychometric function fitted to the pooled data is shown in
the main figures; however, all analyses are based on
psychometric functions fitted to each participant’s data—the
group average PSEs of these functions are shown in the
insets. In the illusion session, PSEs indicate local-driven
TAE regardless of time of perceived uniformity (except for
condition CX in the lower half group, which shows no
noticeable TAE overall). In the control session, the TAE
shifts to global-driven for longer presented physical
uniformity. (c and d) The correlation between each
participant’s average time (%) of uniformity and dPSE during
the illusion (c) and control (d) session is presented. For
(c), the relationship is established with time of perceived
uniformity, and for (d), the relationship is established
with time of physical uniformity. Only for the latter (d) do
we observe the predicted positive correlation with time,
indicative of a shift towards more global TAE.
PSE = point of subjective equality; TAE = tilt
aftereffect.
TAE by time of uniformity. Physical, but not
perceived uniformity, causes a shift towards global-driven
TAE in a time-dependent manner. (a and b) Participants are
classified into two groups according to whether their
average time of uniformity is below (lower half) or above
(upper half) the sample’s median and depict each group’s
average responses by adapting condition in the illusion (a)
and control (b) session. For illustration purposes, a
psychometric function fitted to the pooled data is shown in
the main figures; however, all analyses are based on
psychometric functions fitted to each participant’s data—the
group average PSEs of these functions are shown in the
insets. In the illusion session, PSEs indicate local-driven
TAE regardless of time of perceived uniformity (except for
condition CX in the lower half group, which shows no
noticeable TAE overall). In the control session, the TAE
shifts to global-driven for longer presented physical
uniformity. (c and d) The correlation between each
participant’s average time (%) of uniformity and dPSE during
the illusion (c) and control (d) session is presented. For
(c), the relationship is established with time of perceived
uniformity, and for (d), the relationship is established
with time of physical uniformity. Only for the latter (d) do
we observe the predicted positive correlation with time,
indicative of a shift towards more global TAE.PSE = point of subjective equality; TAE = tilt
aftereffect.Therefore, evidence opposed a positive association between time of
perceived uniformity and a trend towards more global-driven TAE, thus
opposing predictions expected for illusion-based adaptation.For the control session, we performed analogous analyses as for the
illusion session, but with time of physical instead of perceived
uniformity.Because global uniformity is a physical stimulus in this session, a
time-dependent shift from local to global-driven TAE should be expected
regardless of the capacity of illusory orientation to induce a TAE.
Thus, this control session acts as a sanity check to rule out that the
failure to find time-dependency in the illusion session was simply due
to insufficient exposure to the global pattern—even in the cases of
longest time of uniformity.We performed a Bayesian bivariate correlation between individual average
time of physical uniformity and dPSE (Figure 3(d)). Pearson’s
correlation coefficient was r = .468 (95% credible
intervals .056, .714), showing moderate evidence for a positive
correlation: BF+0 = 5.546.Thus, physical uniformity presented for durations equivalent to the
reported illusory uniformity was sufficient to observe a shift towards a
global-driven TAE.
Discussion
The UI is a striking phenomenon in which experience across the whole visual field is
modified by higher precision foveal information, yet its underlying mechanisms
remain unknown. Using a version of UI with oriented Gabor patches, we found that UI
does not produce an orientation adaptation aftereffect consistent with the illusory
percept. Instead, orientation aftereffects only ever followed the (local) physically
presented orientation. This suggests that the UI, at least in orientation, arises
from higher level (higher than primary visual cortex) perceptual processes.It has been suggested that the UI may result from predictive processing operations in
the visual hierarchy (Otten
et al., 2016). In a hierarchical predictive coding scheme, perception
arises from the interaction of bottom-up sensory signals with top-down expectations
generated in higher cortical areas (Friston, 2005; Rao & Ballard, 1999). Prediction error
is determined by the discrepancy between bottom-up sensory signals and the top-down
predictions and propagates through the sensory hierarchy to update the internal
world model. Although the interplay between neural signatures of sensory adaptation
and predictive coding is not fully understood (Symonds et al., 2017), evidence indicates
that top-down expectations produce activity changes in the visual cortex also
specifically for orientation-selective neurons in V1 (Schummers, Sharma, & Sur, 2005), with
adaptation adjusting the relative weight of bottom-up and top-down signals in
relation to their precision (Malmierca, Anderson, & Antunes, 2015). Under this framework, UI may
be conceptualized as the result of high-precision foveal signals being given more
weight in forming perceptual predictions for the presented pattern, possibly in
combination with a prior for perceptual uniformity for the entirety of the visual
field. After a period of exposure, adaptation renders low-precision peripheral
signals weaker still, until eventually they become unable to overcome the
central-based prediction (Otten
et al., 2016).Our results suggest that if UI does result from such predictive operations, the locus
of influence of the feedback does not reach primary visual cortex, as illusory
uniformity produced no measurable adaptation effect.What, then, is the neural basis of UI? UI might be an instance of perceptual
filling-in, a phenomenon whereby a visual attribute like colour, luminance, or
texture is perceived in a region of the visual field even though it only exists in
the surround (Komatsu,
2006). However, unlike typical instances of uniform spread of colour or
luminance, in our orientation UI, the distinction between background and grid
elements persists, and the illusion selectively informs the appearance of the
individual Gabors. The process may be similar to texture filling-in or involve
texture processing in a broader sense. Notably, several neurophysiological and
neuroimaging experiments have reported changes in neural activation in early visual
areas that correlate with perceptual filling-in; however, while for colour or
luminance, this correlate has been seen at V1 (Hsieh & Tse, 2010), for texture
filling-in, it has only been observed at V2 and above (De Weerd, Gattass, Desimone, & Ungerleider,
1995; Komatsu,
2006), in agreement with our results.UI also exhibits similarities with crowding, as a context-dependent alteration of
peripheral perception. Like UI, crowding arises for different low- and high-level
dimensions and at several stages of the visual system, involving V2 and above (Whitney & Levi, 2011),
for instance, tilt adaptation to the veridical orientation is present for crowded,
indistinguishable stimuli (He,
Cavanagh, & Intriligator, 1996). Crowding has been likened to texture
perception (Parkes, Lund,
Angelucci, Solomon, & Morgan, 2001). However, as a fundamental
difference with crowding, in UI, peripheral phenomenology is not a mixture of
adjacent stimuli, but the replacement of peripheral appearance by the traits of
sometimes distant foveal elements.Finally, UI may be due to perceptual inflation, whereby apparent detail in the
periphery is not sustained on perceptual content, but due to decisional or
metacognitive biases (Odegaard,
Chang, Lau, & Cheung, 2018). During the control session in our
experiment, where a physically uniform pattern was presented at times, participants
were less prone to report UI during presentation of the nonuniform pattern compared
with the illusion session: This suggests a shift in decision criterion for
uniformity. Importantly, these processes are not exclusive: Possibly both texture
processing and perceptual inflation contribute to UI. Further studies may elucidate
the precise contribution of the different perceptual mechanisms that underlie
foveal-peripheral integration, as demonstrated by UI, and that are central to
naturalistic visual experience. However, our results clearly demonstrate that, at
least for orientation, these mechanisms do not alter neural coding at the primary
visual cortex.Click here for additional data file.Supplemental material for The Illusion of Uniformity Does Not Depend on the
Primary Visual Cortex: Evidence From Sensory Adaptation by Marta Suárez-Pinilla,
Anil K. Seth and Warrick Roseboom in i-Perception