Martin Lages1, Stephanie C Boyle2, Rob Jenkins3. 1. 1 School of Psychology, University of Glasgow. 2. 2 Institute of Neuroscience and Psychology, University of Glasgow. 3. 3 Department of Psychology, University of York.
Abstract
Visual performance of human observers depends not only on the optics of the eye and early sensory encoding but also on subsequent cortical processing and representations. In two experiments, we demonstrated that motion adaptation can enhance as well as impair visual acuity. Observers who experienced an expanding motion aftereffect exhibited improved letter recognition, whereas observers who experienced a contracting motion aftereffect showed impaired letter recognition. We conclude that illusory enlargement and shrinkage of a visual stimulus can modulate visual acuity.
Visual performance of human observers depends not only on the optics of the eye and early sensory encoding but also on subsequent cortical processing and representations. In two experiments, we demonstrated that motion adaptation can enhance as well as impair visual acuity. Observers who experienced an expanding motion aftereffect exhibited improved letter recognition, whereas observers who experienced a contracting motion aftereffect showed impaired letter recognition. We conclude that illusory enlargement and shrinkage of a visual stimulus can modulate visual acuity.
Visual acuity is commonly assessed using a logarithm-of-the-minimum-angle-of-resolution
(LogMAR) chart (Bailey & Lovie,
1976; Ferris, Kassoff,
Bresnick, & Bailey, 1982). The chart contains letters in unpredictable
sequences; these sequences are arranged in horizontal rows, and the rows are stacked
vertically. Letters in successive rows are printed at smaller sizes, so that they are
increasingly difficult to read. The observer’s task is to read as far down the chart as
possible while maintaining a fixed viewing distance. Visual acuity is calculated from
the size at which the observer can no longer identify the letters reliably (Arditi & Cagenello, 1993;
Snellen, 1862).This method is based on a simple principle: Visual stimuli are easier to resolve when
they are large than when they are small, because a larger image carries more detail.
This principle is uncontroversial for physical changes in image size, for example, when
letter size increases or viewing distance decreases. However, neuroimaging studies have
revealed parallels between apparent size and physical size at the level of cortical
representation. In particular, the perceived size of a stimulus can modulate the spatial
extent of its representation in V1, even when physical stimulus size is held constant
(Fang, Boyaci, Kersten, &
Murray, 2008; Murray,
Boyaci, & Kersten, 2006; Ni, Murray, & Horwitz, 2014; Sperandio, Chouinard, & Goodale,
2012).One method of altering perceived stimulus size while maintaining physical stimulus size
is through prolonged adaptation to spiral motion (Holland, 1965; Thompson, 1880). If an observer adapts to
contracting or expanding motion of a rotating spiral, a subsequent static image appears
to expand or contract in the opposite direction around fixation (Lages, Adams, & Graf, 2009). This
well-established phenomenon is known as the spiral motion aftereffect—a
specific version of the motion aftereffect (Addams, 1834; Anstis, Verstraten, & Mather, 1998). The
spiral motion aftereffect allows one to study consequences of perceived image size
without having to manipulate physical image size or viewing distance (Kersten & Murray, 2010;
Schindel & Arnold,
2010; Sloan,
1951). For example, it allows one to ask whether illusory enlargement of letters
can enhance their identification.In two experiments, we assessed the impact of perceived stimulus size on visual acuity
after motion adaptation. In Experiments 1a and 1b, we employed letters with different
but fixed font sizes in a between-subjects design, and in Experiment 2, we used adaptive
font sizes in a within-subjects design. Our results demonstrate that the motion
aftereffect can modulate visual acuity. More specifically, illusory expanding motion can
improve visual acuity, while illusory contracting motion can decrease visual acuity.
General Method
Both of the experiments adhered to the guidelines of the Declaration of Helsinki. An
initial power analysis suggested that a sample size between 30 and 46 observers
would achieve the desired effect size (Cohen’s f = 0.20–0.25, α =
.05, β = 0.90). Participants were students from the University of Glasgow between
the ages of 16 and 28 years. Only participants who had normal or corrected-to-normal
visual acuity of 10/10 (0.0 logMAR) on a logarithmic visual-acuity chart (Catalog
No. 2103, Precision Vision, Woodstock, IL) took part in the experiments.The stimulus and task were programmed in MATLAB (The MathWorks, Natick, MA) using the
Psychophysics Toolbox extensions (Brainard, 1997; Pelli, 1997). In both experiments, we used
Helvetica bold capital as the text font for the Sloan letters C, D, H, K, N, O, R,
S, V, and Z (Sloan,
1951). These letters were randomly selected and anti-aliased. Letters were
displayed on screen at a viewing distance of 3 m (10 feet) in a dark room with the
lights switched off.Each observer completed a preadaptation and an adaptation block. In the preadaptation
block, a test image with five horizontally aligned letters was presented for 6 s,
and the observer was asked to read out each letter from left to right. An
experimenter recorded each response (whether correct or incorrect). The five letters
were separated horizontally by double spaces and surrounded by a circular Voronoi
pattern on a white background (Fig.
1b). The Voronoi pattern consisted of randomly oriented lines that
elicited a strong contracting or expanding motion aftereffect in the test stimulus
after motion adaptation.
Fig. 1.
Illustration of (a) a linear spiral with high-contrast segments, a moving
version of which was presented in the adaptation phase, and (b) an example
test stimulus consisting of five Sloan letters surrounded by a Voronoi
pattern. In the adaptation phase, spirals rotated either clockwise (inducing
adaptation to contracting motion) or counterclockwise (inducing adaptation
to expanding motion). Sloan letters ranged in size from −0.3 to 0.0 on the
logarithm-of-the-minimum-angle-of-resolution (logMAR) chart (Snellen
fraction: 10/5–10/10) and were surrounded by a Voronoi pattern.
Illustration of (a) a linear spiral with high-contrast segments, a moving
version of which was presented in the adaptation phase, and (b) an example
test stimulus consisting of five Sloan letters surrounded by a Voronoi
pattern. In the adaptation phase, spirals rotated either clockwise (inducing
adaptation to contracting motion) or counterclockwise (inducing adaptation
to expanding motion). Sloan letters ranged in size from −0.3 to 0.0 on the
logarithm-of-the-minimum-angle-of-resolution (logMAR) chart (Snellen
fraction: 10/5–10/10) and were surrounded by a Voronoi pattern.In the adaptation block, the same test stimuli were used, but each test stimulus was
preceded by a 30-s motion-adaptation phase. During motion adaptation, the observer
fixated on the center of a black and white linear spiral (see Fig. 1a) that rotated clockwise or
counterclockwise at a speed of 90° per second. Depending on the direction of
rotation, observers adapted to contracting or expanding motion so that the ensuing
motion aftereffect created illusory expansion or contraction in the test images.
Experiments 1a and 1b
Method
In Experiments 1a and 1b, we recruited 45 and 35 participants, respectively. One
participant in Experiment 1a and 5 participants in Experiment 1b had to be
excluded because they did not meet our requirement of 10/10 visual acuity or
were tested under different lighting conditions from the other participants.
This left data from 44 observers (16 male, 28 female) in Experiment 1a and 30
observers (11 male, 19 female) in Experiment 1b for analysis. Each observer
completed a preadaptation and an adaptation block of 16 trials each. A block
featured four sets of trials with letters in different font sizes. The largest
font size was used in the first set of trials, followed by increasingly smaller
font sizes in the next three sets. Five letters of the same font size were
presented in each of the four trials of a set. Thus, an observer had to identify
a total of 80 letters (5 letters × 4 sets × 4 trials) per block, with 20 letters
at each font size. The four font sizes subtended 4.9, 4.0, 3.1, and 2.2 minutes
of arc (arcmin), respectively. Recognizing 4 out of 5 letters at a given size
approximated Snellen fractions of 10/10 (0.0 logMAR), 10/8 (−0.1 logMAR), 10/6.3
(−0.2 logMAR), and 10/5 (−0.3 logMAR), respectively. Adaptation and test stimuli
subtended 2.2 degrees of visual angle on a 21-in. CRT monitor (Vision Master Pro
514, Iiyama, Tokyo, Japan) with a resolution of 0.32 mm per pixel at a refresh
rate of 120 Hz.In Experiment 1a, we sought to determine whether adaptation to contracting motion
followed by an expanding motion aftereffect would improve visual acuity. We
measured the number of letters that the 44 observers correctly recognized at the
four different font sizes, first in the preadaptation block of trials and again
after adaptation to contracting motion in the second block of trials. As
expected, all observers passed the acuity test for the largest letter size (0.0
logMAR) in the preadaptation block, achieving a Snellen fraction of 10/10 or
better.Conversely, in Experiment 1b, we sought to determine whether adapting to
expanding motion followed by a contracting motion aftereffect would impair
visual acuity. As in Experiment 1a, we measured the number of correctly
recognized letters at four different font sizes, first in a preadaptation block
and again after adaptation to expanding motion in a second block. All 30
observers achieved a visual acuity of at least 10/10 in the preadaptation
block.
Results
First, we conducted separate analyses of variance (ANOVAs) on the number of
correct letter recognitions in Experiments 1a and 1b, respectively. The
within-subjects factors were adaptation (preadaptation, adaptation to
contracting motion in Experiment 1a; preadaptation, adaptation to expanding
motion in Experiment 1b), font size (−0.3, −0.2, −0.1, 0.0 logMAR), and letter
position (1–5 from left to right). In order to compare performance change at
different levels of baseline acuity, we introduced a between-subjects factor by
dividing observers into two groups on the basis of their preadaptation
performance: one with normal visual acuity and one with high visual acuity.Figure 2a illustrates
observers’ performance in Experiment 1a plotted against font size. We used a
median split to categorize observers as having either normal or high visual
acuity. We observed a statistically significant main effect between the two
groups, F(1, 42) = 30.8, p < .001, η2 = .42, which confirmed contrasting levels of baseline
performance for observers below and above the sample median. There was also a
main effect of font size, F(3, 126) = 1,705.7,
p < .001, η2 = .98, which reflects the steep increase in performance with
font size.
Fig. 2.
Results of (a) Experiment 1a and (b) Experiment 1b: mean number of
correctly identified letters as a function of font size, separately for
each combination of block and performance group (normal vs. high, based
on a median split). Observers first completed a preadaptation (PA)
block, followed by a block in which they either adapted to contracting
motion (AC; Experiment 1a) or adapted to expanding motion (AE;
Experiment 1b). Error bars denote ±1 SEM, and in some
cases are smaller than the data points. logMAR = logarithm of the
minimum angle of resolution.
Results of (a) Experiment 1a and (b) Experiment 1b: mean number of
correctly identified letters as a function of font size, separately for
each combination of block and performance group (normal vs. high, based
on a median split). Observers first completed a preadaptation (PA)
block, followed by a block in which they either adapted to contracting
motion (AC; Experiment 1a) or adapted to expanding motion (AE;
Experiment 1b). Error bars denote ±1 SEM, and in some
cases are smaller than the data points. logMAR = logarithm of the
minimum angle of resolution.The ANOVA on letter recognition in Experiment 1b also revealed a statistically
significant effect between groups, F(1, 28) = 17.6,
p < .001, η2 = .39, and font size, F(3, 126) = 481.7,
p < .001, η2 = .95 (Fig.
2b); this confirms that there were different levels of performance
between observers with normal and high baseline acuity and an increasing
difficulty to recognize smaller letters.Critically for our hypothesis, results showed that adaptation to contracting
motion in Experiment 1a resulted in statistically significant improvements,
F(1, 42) = 7.80, p = .0078, η2 = .16, when comparing performance in the preadaptation and
adaptation blocks. Significant interactions between adaptation and group,
F(1, 42) = 12.73, p = .001, and among
adaptation, font size, and group, F(3, 126) = 6.98,
p = .001, revealed that the adaptation effect in Experiment
1a was due to observers with normal visual acuity who identified on average 11%
(2.2 out of 20) more letters at −0.2 logMAR and also showed small improvements
at −0.3 and −0.1 logMAR. Observers with high visual acuity showed no significant
improvement for any of the font sizes (Fig. 2a).Importantly, adaptation to expanding motion in Experiment 1b had the opposite
effect: We found a small but statistically significant reduction in performance,
F(1, 28) = 8.01, p = .0085, η2 = .22, when comparing the preadaptation and adaptation
blocks. This time, however, there were no significant interactions qualifying
the adaptation effect. On average, observers with high visual acuity recognized
7.5% (1.5 out of 20) fewer letters at −0.3 logMAR after motion adaptation, and
observers with normal visual acuity recognized an average of 5% (1.0 out of 20)
fewer letters at −0.1 logMAR (Fig. 2b).We summarized performance across different font sizes by estimating individual
thresholds for letter recognition. Separately for each experiment, we fitted
cumulative Gaussian functions (Wichmann & Hill, 2001) to the
individual data from each block, derived the 50% thresholds for the
preadaptation and adaptation block, and computed the difference between those
thresholds. Comparing the threshold differences in each group revealed a
statistically significant change in performance (M = −0.41
arcmin, 95% confidence interval = [–0.61, –0.21]), two-samples
t(63) = −4.1, p = .0001.Figure 3 illustrates the
significant main effects of letter position in Experiment 1a,
F(4, 168) = 9.42, p < .0001, η2 = .18, and in Experiment 1b, F(4, 112) =
9.86, p < .001, η2 = .26. The letter-position effect indicates that letters at
the center of the display attracted significantly more errors compared with
letters on the left and right. The significant effects of letter position
suggest a form of foveal crowding in which flanking letters impair recognition
of letters at the center (Lev, Yehezkel, & Polat, 2014; Levi, Klein, & Aitsebaomo, 1985).
However, we found no significant interaction between adaptation and letter
position in Experiments 1a and 1b (Fs < 1). This indicated
that crowding and illusory motion affected letter recognition independently of
each other.
Fig. 3.
Results of (a) Experiment 1a and (b) Experiment 1b: mean error as a
function of letter position and block. Mean error was defined as the
number of unrecognized letters averaged across font sizes and observers.
Observers completed two blocks: preadaptation (PA) and either adaptation
to contracting motion (AC; Experiment 1a) or adaptation to expanding
motion (AE; Experiment 1b). Error bars denote ±1
SEM.
Results of (a) Experiment 1a and (b) Experiment 1b: mean error as a
function of letter position and block. Mean error was defined as the
number of unrecognized letters averaged across font sizes and observers.
Observers completed two blocks: preadaptation (PA) and either adaptation
to contracting motion (AC; Experiment 1a) or adaptation to expanding
motion (AE; Experiment 1b). Error bars denote ±1
SEM.In Figures 4a and 4b, visual acuity in
preadaptation is plotted against performance change for each observer in
Experiment 1a and Experiment 1b, respectively. Performance change is expressed
as the difference between correctly identified letters in the adaptation and
preadaptation conditions. Figure 4a shows a negative association between initial visual acuity
and performance change resulting from the expanding motion aftereffect
(Pearson’s r = −0.81, p < .001;
R2 = .658). Together with Figure 2a, this suggests that observers
with high visual acuity did not benefit from the expanding motion aftereffect,
whereas observers with normal visual acuity showed improved acuity at font sizes
−0.1 and −0.2 logMAR.
Fig. 4.
Results of (a) Experiment 1a and (b) Experiment 1b: scatterplots (with
best-fitting regression lines) showing the relationship between
preadaptation (PA) performance and change in performance between the PA
block and the adaptation block. Observers adapted to either contracting
motion (AC; Experiment 1a) or expanding motion (AE; Experiment 1b). In
both plots, PA performance is expressed as both the mean number and the
proportion of correctly identified letters. The vertical line splits
observers into groups with normal and high PA visual acuity. The
horizontal line separates observers with improved performance from
observers with impaired performance.
Results of (a) Experiment 1a and (b) Experiment 1b: scatterplots (with
best-fitting regression lines) showing the relationship between
preadaptation (PA) performance and change in performance between the PA
block and the adaptation block. Observers adapted to either contracting
motion (AC; Experiment 1a) or expanding motion (AE; Experiment 1b). In
both plots, PA performance is expressed as both the mean number and the
proportion of correctly identified letters. The vertical line splits
observers into groups with normal and high PA visual acuity. The
horizontal line separates observers with improved performance from
observers with impaired performance.In contrast to Experiment 1a, there was reduced visual acuity as a consequence of
the contracting motion aftereffect in Experiment 1b and no significant
association between initial acuity and performance change (Fig. 4b; Pearson’s r =
−.039, p = .838; R2 = .0015). In an
additional ANOVA with mixed effects, we combined the data from Experiments 1a
and 1b (N = 74) and entered change in performance (adaptation
block – preadaptation block) as the dependent variable. Font size and letter
position were within-subjects factors, and experiment (1a, 1b) was a
between-subjects factor. The results confirmed that adaptation to contracting
and expanding motion produced the only statistically significant effect on
change in performance, F(1, 72) = 15.1, p <
.001, η2 = .17.
Experiment 2
In a follow-up experiment, we varied motion adaptation within subjects in
separate sessions over consecutive days. A total of 41 naive students from the
University of Glasgow were recruited, but 3 students did not meet the
requirement of 10/10 visual acuity, and a further 6 did not complete the second
session. As a result, data of 32 observers (age range = 19–23 years; 12 male, 20
female) were available for analysis. Instead of fixed font sizes, we employed an
adaptive staircase method to determine individual thresholds of letter size
(Watson & Pelli,
1983). As a consequence, letter size varied from trial to trial
depending on previous responses and letter size. Stimuli were displayed on a
high-resolution 27-in. LED monitor (Cinema Display, Apple, Cupertino, CA) with a
resolution of 0.23 mm per pixel at a refresh rate of 60 Hz and mean luminance of
89 cd/m2.Each observer attended two sessions on consecutive days at the same time of day.
Each session consisted of a preadaptation and an adaptation block of 40 trials
each. Observers adapted to one form of motion in the first session and the other
form of motion in the following session; the sequence of the adaptation
conditions (adaptation to contracting motion first, adaptation to expanding
motion first) was counterbalanced across participants to control for possible
carryover effects.The differences between threshold estimates from the preadaptation and adaptation
blocks on each day were entered into a two-way ANOVA with session (Session 1,
Session 2) and sequence (adaptation to contracting motion first, adaptation to
expanding motion first) as factors. A negative threshold difference indicated
improved performance, and a positive threshold difference indicated impaired
individual performance. The analysis produced a statistically significant
interaction between session and sequence, F(1, 30) = 5.22,
p = .03, η2 = .15.The interaction shown in Figure
5a illustrates the expected opposite adaptation effects for the two
sequences: The average threshold difference for adaptation to contracting motion
in Session 1 is lower compared with adaptation to expanding motion in Session 2
for observers who adapted to contracting motion prior to adapting to expanding
motion. Similarly, the average threshold difference for adaptation to expanding
motion in Session 1 was higher compared with adaptation to contracting motion in
Session 2 for observers who adapted to expanding motion in the first session. In
Figure 5b,
thresholds of preadaptation are plotted against threshold differences between
adaptation and preadaptation blocks for each observer. Despite a sizeable
overlap between the two adaptation conditions, there was a statistically
significant change in performance between adaptation to expanding motion and
contracting motion (M = −0.20 arcmin, 95% confidence interval =
[–0.37, –0.03]); one-sample t(31) = −2.28, p =
.029.
Fig. 5.
Results of Experiment 2. The graph (a) shows the difference in
letter-recognition thresholds between the preadaptation (PA) and
adaptation blocks at each session, separately for observers who adapted
to contracting motion (AC) first and observers who adapted to expanding
motion (AE) first. Error bars denote ±1 SEM. The
scatterplot (b; with best-fitting regression line) shows the
relationship between the PA letter-recognition threshold and the
threshold difference between the PA and adaptation blocks, separately
for observers in the AC and AE groups. The vertical line splits
observers into groups with normal and high PA visual acuity. The
horizontal line separates observers with improved performance from
observers with impaired performance.
Results of Experiment 2. The graph (a) shows the difference in
letter-recognition thresholds between the preadaptation (PA) and
adaptation blocks at each session, separately for observers who adapted
to contracting motion (AC) first and observers who adapted to expanding
motion (AE) first. Error bars denote ±1 SEM. The
scatterplot (b; with best-fitting regression line) shows the
relationship between the PA letter-recognition threshold and the
threshold difference between the PA and adaptation blocks, separately
for observers in the AC and AE groups. The vertical line splits
observers into groups with normal and high PA visual acuity. The
horizontal line separates observers with improved performance from
observers with impaired performance.
Discussion
The present findings convincingly demonstrate that visual acuity can be modulated by
previous adaptation to contracting or expanding motion. In Experiment 1a, the
expanding motion aftereffect improved letter recognition in observers with normal
visual acuity, whereas observers with high visual acuity performed at ceiling and
could not improve further. In Experiment 2, individual observers adapted to
contracting as well as expanding motion in separate sessions. Using an adaptive
staircase method in a within-subjects design, we confirmed the differential effect
of motion adaptation on letter recognition. In particular, illusory expansion in the
test stimulus helped identification of smaller letters that were difficult to
discern otherwise.What is the most plausible explanation for the effect of motion adaptation on visual
acuity? We can rule out pupil size as a confounding factor (Kloosterman et al., 2015; Laeng & Endestad,
2012), since the test stimuli were considerably brighter than the adaptation
stimulus: Luminance of the test stimulus measured on screen changed by 41
cd/m2 in Experiment 1 and by 92 cd/m2 in Experiment 2. In
a control study (N = 6), we confirmed that all observers’ pupils
contracted immediately after onset of the test stimulus in all conditions, but there
was no opposite effect on pupil size during the expanding and contracting motion
aftereffect. Other possible explanations, such as adaptation to blur (Mon-Williams, Tresilian, Strang,
Kochhar, & Wann, 1998; Pseudovs & Brennan, 1993; Webster, Georgeson, & Webster,
2002) and motion blur (Barlow & Olshausen, 2004) did not predict opposite effects on
performance as found in Experiments 1 and 2.It is possible that the motion aftereffect shifted and scaled the representation of
the stimulus in cortical area V1 (Murray et al., 2006; Ni et al., 2014; Sperandio et al., 2012; Whitney et al., 2003). More
specifically, the motion aftereffect may have modulated surround suppression in the
retinotopically organized receptive fields of the medial temporal brain region
(Anton-Erxleben, Stephan,
& Treue, 2009), which possibly altered spatial resolution at earlier
processing stages through recurrent feedback (Anton-Erxleben & Carrasco, 2013; Carrasco, Williams, &
Yeshurun, 2002; Fang
et al., 2008). Although this explanation is rather speculative (e.g.,
Morgan, 2012), a more
extensive neural representation in retinotopically organized cortical areas would
make it easier to resolve discriminating features during object recognition (Barlow, 1961; Vinje & Gallant,
2000).Another possible explanation is that the expanding motion aftereffect did not alter
the neural representation but facilitated the “readout” or recognition of letters.
The contracting motion aftereffect may have increased foveal crowding, whereas the
expanding motion aftereffect may have released crowding between letters and the
surrounding Voronoi pattern (Lev
et al., 2014; Levi et
al., 1985). This would help and impede recognition of letters,
respectively (Herzog, Sayim,
Chicherov, & Manassi, 2015). Although the position effect in
Experiments 1a and 1b is reminiscent of the effect of crowding between letters in
the periphery (Bouma,
1970; Dakin, Greenwood,
Carlson, & Bex, 2011; Falkenberg, Rubin, & Bex, 2007; Pelli & Tillman, 2008;
Whitney & Levi,
2011), we found no significant interaction between adaptation and letter
position. This suggests that any crowding effect was independent of the motion
aftereffect.We conclude that an illusory change in perceived stimulus size after motion
adaptation is responsible for this small but surprising effect. Visual acuity is
routinely associated with the optics of the human eye and refractive errors. The
striking implication of our findings is that, under the conditions described here,
adaptation to contracting motion can improve visual acuity.
Authors: David Whitney; Herbert C Goltz; Christopher G Thomas; Joseph S Gati; Ravi S Menon; Melvyn A Goodale Journal: Science Date: 2003-09-18 Impact factor: 47.728