Eye Movements, Strabismus, Amblyopia and Neuro-Ophthalmology

Adaptation to Laterally Displacing Prisms in

Anisometropic Amblyopia
Jaime C. Sklar,1 Herbert C. Goltz,1,2 Luke Gane,1 and Agnes M. F. Wong1–3
1
Neurosciences and Mental Health, The Hospital for Sick Children, Toronto, Canada
2
Department of Ophthalmology and Vision Sciences, University of Toronto, Toronto, Canada
3Department of Ophthalmology and Vision Sciences, The Hospital for Sick Children, Toronto, Canada

Correspondence: Agnes M. F. Wong, PURPOSE. Using visual feedback to modify sensorimotor output in response to changes in the
Department of Ophthalmology and external environment is essential for daily function. Prism adaptation is a well-established
Vision Sciences, The Hospital for experimental paradigm to quantify sensorimotor adaptation; that is, how the sensorimotor
Sick Children, 555 University Ave- system adapts to an optically-altered visuospatial environment. Amblyopia is a neurodevel-
nue, Toronto, ON M5G 1X8, Canada;
[email protected].
opmental disorder characterized by spatiotemporal deficits in vision that impacts manual and
oculomotor function. This study explored the effects of anisometropic amblyopia on prism
Submitted: February 3, 2015 adaptation.
Accepted: April 22, 2015
METHODS. Eight participants with anisometropic amblyopia and 11 visually-normal adults, all
Citation: Sklar JC, Goltz HC, Gane L,
Wong AMF. Adaptation to laterally
right-handed, were tested. Participants pointed to visual targets and were presented with
displacing prisms in anisometroic feedback of hand position near the terminus of limb movement in three blocks: baseline,
amblyopia. Invest Ophthalmol Vis Sci. adaptation, and deadaptation. Adaptation was induced by viewing with binocular 11.48 (20
2015;56:3699–3708. DOI:10.1167/ prism diopter [PD]) left-shifting prisms. All tasks were performed during binocular viewing.
iovs.15-16605 RESULTS. Participants with anisometropic amblyopia required significantly more trials (i.e.,
increased time constant) to adapt to prismatic optical displacement than visually-normal
controls. During the rapid error correction phase of adaptation, people with anisometropic
amblyopia also exhibited greater variance in motor output than visually-normal controls.
CONCLUSIONS. Amblyopia impacts on the ability to adapt the sensorimotor system to an
optically-displaced visual environment. The increased time constant and greater variance in
motor output during the rapid error correction phase of adaptation may indicate deficits in
processing of visual information as a result of degraded spatiotemporal vision in amblyopia.
Keywords: prism adaptation, amblyopia, anisometropia, manual pointing, sensorimotor
integration

mblyopia, or ‘‘lazy-eye,’’ is a neurodevelopmental visual tasks in everyday life,28 especially ones that require visuomotor
A disorder that results from abnormal visual stimulation
during early childhood, usually caused by anisometropia
integration to optimize performance.29,30 Because vision
normally provides the most reliable sensory signals when
(interocular difference in refractive errors), strabismus (eye detecting subtle changes in the environment, it often is used to
misalignment), or a mix of the two.1–4 It generally is defined as calibrate and guide motor actions.31–33 To the best of our
a unilateral (or less often bilateral) decrease in visual acuity not knowledge, our group was the first to investigate the ability of
directly attributable to a structural pathology of the eye itself the sensorimotor system in amblyopia to adapt to an
and cannot be corrected immediately by optical means.5,6 experimental perturbation in the visual environment24 by
The sensory deficits associated with amblyopia have been using a double-step saccade adaptation paradigm.34,35 During
researched extensively. For example, people with amblyopia short-term gain-down saccadic adaptation, participants with
exhibit deficits in spatial localization as a result of positional anisometropic amblyopia exhibited an impaired spatial re-
uncertainty,4,7–9 increased spatial and temporal crowding,10,11 sponse to saccadic adaptation, as evidenced by diminished
and deficits in global motion perception.12–14 These deficits saccadic gain modulation.24 In a follow-up experiment,36 we
have been attributed to increased noise in the amblyopic visual demonstrated that a similar decrease in saccadic gain could be
system,15–17 spatial undersampling of visual neurons in the replicated by introducing an equivalent amount of exogenous
striate cortex,18–20 and spatial distortions.21,22 Visuomotor spatial noise to the adapting step in normal participants,
integration in amblyopia, however, has not been examined as providing strong support for the hypothesis that the reduction
extensively. We have documented previously visuomotor in saccadic adaptation in amblyopia was the result of a less
deficits in amblyopia, including saccades with longer latency precise visual error signal driving adaptation.
and less precision,23,24 altered temporal dynamics of visually- The finding that amblyopia alters oculomotor adaptation
guided reach-to-touch movements,25,26 and impaired online led us to ask whether visuomanual adaptation (e.g., prism
motor control, especially in severe amblyopia.27 adaptation) also would be impacted. Prism adaptation is a
The ability to adapt or modify behavior in response to a well-established experimental method for the study of
changing external visual environment is essential to completing sensorimotor adaptation.37–45 After shifting the visual world

Copyright 2015 The Association for Research in Vision and Ophthalmology, Inc.
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TABLE. Clinical Characteristics for Participants With Anisometropic Amblyopia

Visual Acuity Refractive

Snellen (logMAR) Correction
Amblyopia Deviation at Stereoacuity,
ID Age Sex Severity Right Eye Left Eye Near, 0.33 m, D Right Eye Left Eye arcsec

1 27 F Mild 20/40 (0.30) 20/20 (0.00) LET1, ETþX4 þ0.25 þ2.75 140
2 46 F Moderate 20/100 (0.70) 20/20 (0.00) RET 2 þ2.25 þ0.25 3 174 0.75 3000
3 26 F Mild 20/20 (0.00) 20/50 (0.40) E4 1.50 þ1.50 3 80 3.00 þ2.50 3 80 120
4 36 F Mild 20/20 (0.00) 20/50 (0.40) LET2 1.50 þ0.50 3 165 Plano 140
5 20 F Mild 20/20 (0.00) 20/60 (0.48) LET2 1.50 þ0.50 3 80 þ1.00 þ1.25 3 95 200
6 18 F Mild 20/20 (0.00) 20/32 (0.20) X1 0.75 þ2.00 3 84 2.75 þ4.50 3 99 40
7 24 M Mild 20/25 (0.10) 20/15 (0.10) RXT2 þ2.50 Plano 3000
8 42 M Moderate 20/100 (0.70) 20/20 (0.00) RET2 9.25 þ0.75 3 90 8.75 þ2.25 3 90 3000

optically using wedge prisms, the sensorimotor system eye deviations (by the prism and cover tests). Exclusion criteria
adjusts to compensate for the new visuospatial environ- included any eye deviations on the cover test of >8 prism
ment.39 For example, when a person points to a visual target diopters (PD) indicating a strabismic or mixed-mechanism
while viewing through wedge prisms that displace the visual amblyopia,49 ocular pathology resulting in reduced visual
world optically in the leftward direction, an initial pointing acuity, previous ocular surgery, or neurologic disease. Written
error to the left of the target occurs. Subsequently, as the consent was obtained to enroll in this study. This investigation
person points repeatedly to the target over several trials in the was approved by the Research Ethics Board at The Hospital for
presence of visual feedback, this pointing error decreases to Sick Children and conformed to the Declaration of Helsinki.
the baseline level.29,39,45 When the prisms are removed,
pointing error in the opposite direction occurs (i.e., to the Apparatus
right of the target position) before performance returns to
baseline after repeated pointing in the presence of visual A large drafting table inclined at 458 was placed 46 cm in front
feedback. This is termed the negative aftereffect.44 Prism of participants at eye level.50 Visual targets generated by a
adaptation consists of two phases: The first phase is a rapid custom Cþþ program were displayed on a cathode ray tube
error correction phase, in which visually normal participants monitor (Mitsubishi CRT Diamond Pro 2070SB-BK, viewing
correct their motor errors iteratively over successive trials. area of 20 3 15 inches; Cambridge Research Systems,
This phase is thought to represent a high-level spatial Rochester, Kent, UK) at 120 Hz and reflected off a semi-
remapping by the cognitive system to reduce performance silvered mirror. These small square white targets subtended a
error quickly.44,45 For instance, performance error can be visual angle of 0.258 at a viewing distance of 46 cm, and were
reduced rapidly by using error information obtained from a presented randomly along a single horizontal axis at either 08,
previous pointing trial to guide the next trial (i.e., resetting 638, or 698 on a virtual plane. The targets were perceived as
the movement plan using feedback at the end of the trial), or being coplanar with the table surface. The horizontal and
by the strategic use of online visual feedback.46–48 The second vertical axes constituted an orthogonal basis for the virtual
phase is a plateau phase in which visually normal participants plane. The horizontal and vertical axes lay in the plane of the
reach the same precision and accuracy as during baseline table, which was inclined 458 relative to the transverse plane of
pointing. This phase is thought to be based on offline the participant. The vertical axis lay in the sagittal plane and
feedback mechanisms.39,48 In this study, we investigated the the horizontal axis was parallel to the participant’s coronal
effects of anisometropic amblyopia on adaption to displace- plane (Fig. 1).
ment of the visual world using wedge prisms, a well- Reaching movements were recorded using an infrared
established experimental paradigm for the study of sensory- motion capture system (Optotrak Certus, spatial accuracy 0.1
motor adaptation. mm, resolution 0.01 mm, sampling frequency 200 Hz;
Northern Digital, Waterloo, Canada). All pointing movements
were made using the dominant right hand of the participant.
METHODS The infrared marker used by the motion capture system to
record finger position throughout the movement trajectory
Participants was secured to the right index finger. Visual feedback of finger
position was presented to the participant in real-time as a 1.28
A total of 11 visually-normal controls (6 males, age ¼ 29 6 8 circle rendered on the CRT and reflected onto the semi-silvered
years) with normal or corrected-to-normal vision, and eight mirror; thus, appearing on the same plane as the visual targets.
participants with anisometropic amblyopia (2 males, age ¼ 32 This feedback was only presented near the terminus of
6 11 years, see Table), all right-handed, participated in this movement, defined here as when the finger position reached
study. All participants were tested with their habitual optical 75% of the elevation-axis distance to the target from a
correction. Amblyopia was defined as visual acuity of 20/25 or consistent start position (indicated by a coin affixed to the
worse in the amblyopic eye (range, 20/25–20/100), 20/20 or surface of the drafting table that served as a tactile cue) in all
better in the fellow eye, and an interocular difference of two or three experimental blocks. Terminal feedback was chosen to
more chart lines. Anisometropic amblyopia was defined as reduce oblique and/or secondary movements,51,52 especially
amblyopia in the presence of an interocular refractive error during prism adaptation trials.
difference of ‡ 1 diopter (D) in spherical or cylindrical power.
All participants underwent a full assessment by a certified Experimental Protocol
orthoptist, who assessed visual acuity (using the Early
Treatment of Diabetic Retinopathy Study [ETDRS] chart), stereo The experimental procedure comprised 3 blocks: baseline (50
acuity (using the Randot test), fusion (using Worth-4-Dot), and trials, Fig. 2A), adaptation (200 trials, Fig. 2B), and dead-

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FIGURE 1. The experimental apparatus used for the prism adaptation protocol. Hand movements were tracked by an Optotrak Certus. Visual
targets were rendered on a CRT monitor and reflected off a semi-silvered mirror such that they appear coincident with the plane of the inclined
drafting table. Hand position feedback appeared in real time as a small white circle when the right index finger position reached 75% of the
elevation axis distance to the target.

aptation (70 trials, Fig. 2C). During all three experimental these lenses was approximately 908, and, thus, encompassed
blocks, participants viewed binocularly while wearing goggle the entire stimulus range, in both the prism and nonprism
frames fitted with hinge-mounted 20 PD (~11.48) left-shifting conditions. During the adaptation block, the prisms were
wedge prisms on top of permanently-affixed plano lenses in swung down in front of the eyes. During the baseline and
front of both eyes. The viewing angle for each eye through deadaptation blocks, only the plano lenses were in front of the

FIGURE 2. (A) Baseline pointing task. (B) Prism adaptation task during the (i) rapid error correction phase and (ii) plateau phase. (C) Prism
deadaptation task during the (i) early phase and (ii) plateau phase. Dotted arrows depict the expected trajectory of the hand. Gray square
represents the veridical position of the target of interest.

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eyes. This ensured that the goggles themselves had no impact of the exponential fit were considered outliers and were
on the effect of prism adaptation. eliminated from analysis.
A force-sensitive resistor (FSR,15 mm diameter; Tekscan, Variable error during adaptation between the two groups
Boston, MA, USA) was used to initiate and end each trial, and was compared by calculating the squared mean of residuals for
was triggered by tapping it with the left index finger. It was the rapid error correction and plateau phases of adaptation.
placed outside the stimulus display area at the bottom left The squared mean of residuals characterizes the variance of
corner of the inclined table. The FSR signal was sent to the data points around the estimated exponential function. The
computer controlling the experimental protocol by the analog- squared mean of residuals was calculated by averaging the
to-digital converter of the ViSaGe stimulus generator system squared residual distances from the exponential function for
(Cambridge Research Systems) via a custom interface board. the number of trials equal to the time constant (rapid error
The end of the trial was indicated when the participant pressed correction phase) of each individual participant (except one
the FSR for the second time. participant with amblyopia who did not exhibit a time constant
Participants were instructed to point as accurately as significantly different from zero), as well as the last five trials of
possible to the visual target while pacing their limb movements the adaptation block (plateau phase) using the following
with a metronome set at 1.33 Hz.45,52–54 This frequency was equations:
chosen to reduce the incidence of oblique/secondary move- D ¼ f ðnÞobserved  f ðnÞpredicted
ments, especially during the prism adaptation task. To ensure
that all participants were able to keep to the beat of the
metronome, practice trials were given to each participant f ðnÞpredicted ¼ fo þ að1  ebn Þ
before the baseline trials were initiated (ranging from 20–30
depending on individual performance). where D is vertical distance of the point in question to the
exponential function, f (n)observed is the constant error in the
Data Analysis azimuthal plane for a given trial, and f (n)predicted is the constant
error of the pointing movement in the azimuthal plane as
The main outcome measures were: (1) constant and variable predicted by the exponential model.
error during baseline pointing; (2) magnitude of adaptation, Statistical analysis was conducted using the R 3.1.1 software
rate of adaptation, and variance of the residuals of pointing package,56 where the ez package was used for the ANOVAs
during the rapid error correction and plateau phases of performed.57 All comparisons used the azimuth-axis position
adaptation; and (3) magnitude and rate of deadaptation. All for analysis, as this was the plane of prism perturbation. For
outcome measures used the azimuth (horizontal) axis pointing baseline constant error and variable error, magnitudes of
position for analysis, as this was the plane of the prism adaptation and deadaptation, and the time course of adaptation
perturbation. (and deadaptation), a 3-way ANOVA was performed with
During baseline pointing, the mean accuracy, or constant Group as the between-subject factor (2 levels; visually-normal
error, was defined as the mean difference between terminal controls and participants with anisometropic amblyopia) and
finger position and veridical target position. This was two within-subject repeated factors of Block (3 levels; Baseline,
calculated within subject by computing the mean constant Adaptation, and Deadaptation) and Target Position (5 levels;
error across all 50 baseline block trials. The precision, or 98, 38, 08, 38, and 98) for constant and variable error. In these
variable error, was defined as the mean standard deviation (SD) analyses, a Greenhouse-Geisser correction was used when the
of the constant error across the baseline block.55 The SD was assumption of sphericity was violated. Additionally, in the
calculated within subject using all 50 baseline block trials. adaptation and deadaptation blocks only the last 10 trials for
The time course (rate) and magnitude of adaptation were each target position were taken into consideration, such that
estimated by fitting an exponential rise to maximum function the initial constant error induced by the wedge prisms did not
to all 200 trials during the adaptation block, where the confound the results. These data then were collapsed across all
pointing error spatial magnitude in the azimuthal plane target positions, and each of the 5 outcome measures were
decreased as a consequence of an increasing number of trials. compared between the control and amblyopia groups using
Each participant’s individual pointing data were fitted using the independent sample Student’s t-tests followed by a false
function: discovery rate (FDR) correction to compensate for multiple
planned comparisons. All significant main effects and interac-
f ðnÞ ¼ f0 þ að1  ebn Þ tions were assessed using post hoc Student’s t-test followed by
an FDR correction to correct for multiple comparisons. All
Where f (n) represents the predicted pointing accuracy on a outcome measures are reported as mean 6 SD, and all P values
given trial n, fo denotes the predicted pointing accuracy at n ¼ are reported as adjusted P values after correction for multiple
0, a represents the change in pointing accuracy from n ¼ 0 comparisons. All negative values indicate a leftward bias.
[f (0)] to n  ‘ ( f ‘), and b denotes the rise/decay constant of The variance of residuals during adaptation was analyzed
adaptation. The value of b1 is considered to be the time constant using a 2-way mixed ANOVA with Group as the between-
of adaptation (the number of trials to reach ~63.2% of total subject factor (2 levels; visually-normal controls and partici-
adaptation) and was used for subsequent analysis. Similarly, the pants with anisometropic amblyopia) and Phase of Adaptation
time course (rate) and magnitude of deadaptation were as the repeated within-subject factor (2 levels; rapid error
estimated by fitting an exponential decay function to all 70 correction and plateau phases).
trials during that block using the same equation. Magnitudes of
adaptation and deadaptation were assessed for each participant
individually by computing the difference between predicted RESULTS
pointing response on trial 1 [f (1)] and the asymptotic value of
Baseline Pointing
the exponential fit. This characterizes the magnitude of change
in constant error from the beginning to the end of the There was no significant interaction between Group and Block
adaptation and deadaptation blocks, respectively. Any trials for either constant error (F[2,34] ¼ 2.7, P ¼ 0.08) or variable
that fell on or outside of the 99% confidence intervals (6 3 SD) error (F[2,34] ¼ 0.1, P ¼ 0.9). This indicates that during baseline

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FIGURE 3. (A) Representative data with exponential fits from a visually-normal control and a participant with anisometropic amblyopia. Dotted
lines represent mean constant error of baseline pointing. (B) Comparison of time constants (number of trials to reach ~63% steady-state adaptation)
between the two groups. Participants with anisometropic amblyopia required significantly more trials to prism adapt than visually-normal controls
(adjusted P ¼ 0.03). Error bars: 61 SEM.

pointing, there was no significant difference between controls adaptation between visually-normal controls (0.158 6 0.848)
and participants with amblyopia for constant error (controls ¼ and participants with amblyopia (0.478 6 0.698; t[17] ¼ 0.89, P
0.368 6 0.68, amblyopia ¼0.088 6 0.68; t[93] ¼2.1, adjusted ¼ 0.38).
P ¼ 0.1) or variable error (controls ¼ 0.878 6 0.38, amblyopia ¼ Time Course of Adaptation. In general, prism adaptation
0.858 6 0.28; t[93] ¼ 0.5, adjusted P ¼ 0.9). in both groups demonstrated a well-characterized temporal
pattern, with an initial rapid error correction phase followed
Prism Adaptation by a gradual error correction phase that eventually plateaued at
a steady state (Fig. 3A). However, the time course differed
Magnitude of Adaptation. There was no significant between groups—participants with amblyopia had a mean
difference in mean magnitude of spatial adaptation when time constant of 17 6 10 trials that was significantly longer
comparing visually-normal controls (9.18 6 2.38) and partici- than the 4 6 3 trials time constant for controls (t[6.5] ¼ 3.6,
pants with anisometropic amblyopia (8.28 6 4.58; t[17] ¼ 0.62, adjusted P ¼ 0.03; Fig. 3B).
adjusted P ¼ 0.82). To address whether the greater time constant is associated
As a secondary analysis, we assessed how close the constant with a reduction in binocularity (stereopsis) that accompanies
error at the end of adaptation and deadaptation came to amblyopia versus amblyopia per se, we performed Pearson’s
constant error at the end of the baseline block. To do this, correlation analyses in participants with amblyopia. We found
adaptation and deadaptation were assessed for each participant no significant correlation between stereoacuity (range, 40–
individually by computing the mean of the last 10 baseline 3000 arc sec) and time constant of adaptation (r ¼ þ0.17, P ¼
trials, and subtracting it from the mean of the last 10 trials of 0.72), nor between visual acuity (range, 20/25–20/100) of the
the adaptation block or of the deadaptation block. By the last amblyopic eye and time constant of adaptation (r ¼þ0.61, P ¼
10 trials of the adaptation and deadaptation blocks, all 0.15).
participants in both groups had reached the plateau phase of Variance of Residuals During Prism Adaptation.
the adaptation and deadaptation. There was no difference in Participants with amblyopia exhibited more variance of
the ability to attain baseline constant error values at the end of residuals (less precision) during early adaptation (the rapid

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FIGURE 4. (A) Mean of residuals around the exponential fit for the first 40 trials of the adaptation block in visually-normal controls and participants
with anisometropic amblyopia. (B) Comparison of the squared mean of residuals (variance) between the two groups during the rapid error
correction and plateau phases of the exponential fit for the adaptation block. During the rapid error correction phase of adaptation, a significantly
higher mean variance of residuals was found in participants with amblyopia compared to controls (adjusted P ¼ 0.004). Additionally, among
participants with amblyopia, there was a significantly greater mean variance of residuals during rapid error correction phase compared to plateau
phase of adaptation (adjusted P ¼ 0.02). Error bars: 61 SEM.

error correction phase) than controls (Fig. 4A). There was a (3.18 6 2.98; t[13] ¼ 0.01, adjusted P ¼ 0.99). Additionally,
Group by Phase of Adaptation interaction (F[1,16] ¼ 14.2, P ¼ there was no difference in the ability to attain baseline constant
0.002), with a significantly higher squared mean of residuals in error levels at the end of deadaptation between controls
participants with amblyopia (2.998 6 1.218) compared to (0.478 6 0.68) and participants with amblyopia (0.258 6
controls (0.888 6 1.028; t[16] ¼ 3.9, adjusted P ¼ 0.004; Fig. 0.68; t[17] ¼ 0.82, P ¼ 0.43).
4B) during the rapid error correction phase of adaptation. Time Course of Deadaptation. Only 9 of 19 participants
During the plateau phase of adaptation, there was no (4 visually-normal controls, 5 participants with amblyopia)
significant difference between controls (1.368 6 1.28) and exhibited an exponential decay that was modeled well by the
participants with amblyopia (1.368 6 1.08; t[16] ¼ 0.015, exponential decay fit. This reduced sample size made it
adjusted P ¼ 0.99). Additionally, among participants with difficult to conduct a meaningful statistical analysis on an
amblyopia, there was a significantly greater squared mean of individual basis. When using group means, however, these data
residuals during the rapid error correction phase (2.998 6 were well modeled by an exponential decay fit (Fig. 5). Based
1.218) compared to the plateau phase (1.368 6 1.08; t[6] ¼ 3.6, on the fits of the grouped mean responses, there was no
adjusted P ¼ 0.02), but no such difference was observed in difference in the time course of deadaptation between controls
controls (rapid error correction, 0.888 6 1.028; plateau, 1.368 (5 trials) and participants with amblyopia (5 trials; Fig. 5).
6 1.28; t[10] ¼ 1.4, adjusted P ¼ 0.25).
Movement Duration
Prism Deadaptation
To verify that there were no between group differences in
Magnitude of Deadaptation. Similar to adaptation, there performance on the pointing task, we used a repeated
was no significant difference in the mean magnitude of spatial measures ANOVA to compare movement duration across
deadaptation between visually-normal controls (visually-normal baseline, adaptation, and deadaptation. There was no signifi-
controls ¼ 3.18 6 1.58) and participants with amblyopia cant main effect of Group (F[1,13] ¼ 3.5, P ¼ 0.09) and Block

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FIGURE 5. Comparison of the time constants during deadaptation for participants with anisometropic amblyopia and visually-normal controls using
mean group data. Error bars: 61 SEM.

(F[1.28, 16.6] ¼ 0.50, P ¼ 0.60), nor was there a significant The ability to integrate sensory and motor information
interaction between Group and Block (F[1.28, 16.6] ¼ 0.44, P ¼ properly follows a developmental trajectory.60 During normal
0.64). This indicates that the movement time was consistent development, repeated exposure to coincident visual-motor
across the two groups and the three blocks—all participants stimuli (e.g., when making a visually-guided action to a target)
were able to keep to the beat of the metronome consistently results in the creation of strong relations between motor
under all experimental conditions. commands and the visual outcome of motor actions, which
allows optimal movement accuracy and precision to an
Effects of Target Position on Accuracy and intended target.59,62,63 This developmental course of visuomo-
tor integration is particularly relevant in the context of
Precision During Baseline Pointing, Adaptation, anisometropic amblyopia, a neurodevelopmental disorder of
and Deadaptation spatiotemporal vision. Amblyopia is associated with decreased
For pointing accuracy, a 3-way ANOVA demonstrated a reliability of the visual signal. We have demonstrated previously
significant main effect of Target Position (F[1.5, 25.2] ¼ 13.7, P that during binocular viewing, participants with anisometropic
< 0.001) with a small difference in pointing accuracy among amblyopia demonstrate greater variability in saccadic latencies
target positions (ranging from 0.768 to 0.258; Supplementary and amplitudes,23 and decreased saccadic adaptation,24 com-
Table S1). A significant main effect of Block (F[2, 34] ¼ 29.7, P < pared to visually-normal controls. We hypothesized that these
0.001) also was found with a small difference in pointing deficits occur secondary to more variable visual information,
accuracy among blocks (ranging from 0.438 to 0.878; rather than deficits in the motor system itself.24,36 During early
Supplementary Table S2). childhood, people with amblyopia likely develop atypical
For pointing precision, a significant main effect of Block visuomotor correlations based upon abnormal visual afferent
was found (F[2,34] ¼ 6.5, P ¼ 0.004) attributable to a small information as a result of increased signal variability,15,19,23,24,64
difference in pointing precision among blocks (ranging from spatial undersampling,16,18 and spatial distortions.21,22 We have
0.808 to 1.08; Supplementary Table S3). documented some of these atypical visuomotor interactions
previously in people with amblyopia, including altered
kinematic strategies during visually-guided reaching (e.g.,
DISCUSSION decreased peak acceleration/velocity and a prolonged acceler-
ation phase). In this study, we hypothesized that people with
We investigated the effects of anisometropic amblyopia on anisometropic amblyopia may require more trials to integrate
adaption to lateral displacement of the visual world using the spatial information as a result of a lower-fidelity visual error
wedge prisms, a well-established experimental paradigm for signal available to aid in realigning the sensory and motor
the study of sensory-motor adaptation. When prisms are used coordinate frames after exposure to a laterally displaced visual
to induce a large sensory prediction error that is novel to environment. This idea of a decreased adaptation/learning rate
visually-normal participants, they introduce an offset between secondary to increased sensory noise conforms well with
the altered visual input and the resultant efferent motor output theories of Bayesian inferences in motor learning — for a
for a given pointing action.58–60 During the rapid error consistent perturbation (in this case, the constant optical
correction phase of adaptation, visual information available displacement), the more sensory noise that is present in the
during a movement is used to modify trajectories within a error command, the longer the learning will take.65,66
single trial29 and on subsequent trials48 to allow for compen- Visual input provides an important signal to the cerebellum,
sation in visuomotor function for an optically displaced which not only has a major role in visually-guided coordination
environment, leading to improved pointing accuracy48,53,61 of ongoing movements,67 but also in prism adaptation as
and, hence, prism adaptation.59,60,62 In this study, we showed evident from lesional studies in human55,68 and nonhuman
that participants with amblyopia exhibited the same magnitude primates,37,42 and from functional imaging studies in hu-
of adaptation as visually-normal controls, but they demonstrat- mans.38,69,70 Patients with cerebellar dysfunction exhibit
ed a longer time constant (i.e., required more trials) and reduced or absent adaptation to a shifted optical world, even
increased variance of residuals during the early rapid error after many repeated trials with visual feedback.37,42,55 There-
correction phase of adaptation. fore, it is reasonable to postulate that as a result of decreased

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fidelity of the visual signal reaching the cerebellum in Acknowledgments

anisometropic amblyopia, prism adaptation will be impacted.
We thank Linda Colpa for performing orthoptic examinations.
This is supported by our finding that more trials were required
(i.e., greater time constant) for participants with amblyopia to Supported by Grant MOP 106663 from the Canadian Institutes of
achieve a comparable magnitude of adaptation to visually- Health Research (CIHR), Leaders Opportunity Fund from the
normal controls. Canada Foundation for Innovation (CFI), the Brandan’s Eye
Research Foundation, the John and Melinda Thompson Endow-
ment Fund in Vision Neurosciences, and the Department of
Increased Variance in Residuals During the Rapid Ophthalmology and Vision Sciences at The Hospital for Sick
Error Correction Phase of Adaptation Children.
Disclosure: J.C. Sklar, None; H.C. Goltz, None; L. Gane, None;
Deficits along the dorsal visual pathway have been identified in
A.M.F. Wong, None
people with amblyopia.13,71–73 The dorsal visual pathway, one
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