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Dissociating perceptual and motor effects of prism
adaptation in neglect
Christopher L. Striemer
a
and James Danckert
b
Prism adaptation reduces some symptoms of neglect;
however the mechanisms underlying such changes are
poorly understood. We suggest that prisms influence
neglect by acting on dorsal stream circuits subserving
visuomotor control, with little influence on perceptual
aspects of neglect. We examined prism adaptation in three
neglect patients and a group of healthy controls on line
bisection and landmark tasks. Neglect patients showed a
dramatic reduction in the rightward bias for line bisection,
but absolutely no change in their leftward bias on the
landmark task, which is a perceptual equivalent to bisection.
However, in controls, prisms produced ‘neglect-like’
deficits on both the line bisection and landmark tasks.
These data suggest that prisms influence visually guided
actions more so than perception in neglect. NeuroReport
21:436–441 c2010 Wolters Kluwer Health | Lippincott
Williams & Wilkins.
NeuroReport 2010, 21:436–441
Keywords: attention, neglect, parietal lobe, perception–action, prism
adaptation
a
Department of Psychology, Centre for Brain and Mind, Social Sciences Centre,
University of Western Ontario, London and
b
Department of Psychology,
University of Waterloo, Waterloo, Ontario, Canada
Correspondence to James Danckert, PhD, Department of Psychology, University
of Waterloo, 200 University Avenue West, Waterloo, Ontario N2L 3G1, Canada
Tel: + 1 519 888 4567 x37014; fax: + 1 519 746 8631;
e-mail: jdancker@uwaterloo.ca
Received 1 February 2010 accepted 3 February 2010
Introduction
Lesions to right parietal cortex often lead to neglect – a
disorder in which patients are unaware of contralesional
space [1]. One method that has been shown to improve
neglect symptoms is prism adaptation (PA) [2], in which
patients point to targets while wearing prisms that shift
vision 101rightward. Before PA, neglect patients’
subjective straight-ahead judgments are biased right of
center. During adaptation patients adjust their pointing
leftward to compensate for the rightward prism shift.
After 5 min, patients’ straight-ahead judgments are shifted
leftward, closer to center. After adaptation, patients
improve on clinical and experimental tasks including tactile
perception, occulomotor biases, and spatial attention (for a
reviewseeRef.[3]).
Although prisms improve some neglect symptoms, the
mechanisms for these effects are unknown. How does
visuomotor adaptation, relying on mechanisms in cere-
bellar, parietal, and motor cortex [4,5], improve such a
broad range of symptoms? The dual pathways’ model
of vision suggests that visuomotor control is subserved
by the dorsal stream (V1 to posterior parietal cortex),
whereas conscious perception is subserved by the
ventral stream (V1 to inferior temporal cortex) [6]. One
explanation for the effects of prisms is that they alter
dorsal stream behaviors – the pathway most active during
adaptation [4]. That is, leftward realignment signals,
generated in the right cerebellum, are sent to left
posterior parietal cortex [3,7]. The realignment signals
encourage the patient to shift exploratory motor beha-
viors leftward into previously neglected space, resulting
in a decrease in some symptoms [3]. This would explain
many of the effects of prisms on tasks requiring overt
motor responses with the adapted limb (e.g., line bisection,
cancellation, figure copying). Furthermore, many of the
‘perceptual’ after effects of prisms such as covert attention
[8] or visual imagery mental imagery [9,10] may be related
to effects of PA on eye movements [11–13], which are
also controlled by the dorsal stream [6]. This could allow
patients to mentally scan a previously learned image [14]
without necessitating any changes in bottom-up visual
perception. Importantly, the dorsal stream is typically
undamaged in neglect [1]. Furthermore, recent imaging
[15] and patient studies [16] implicate the dorsal stream as
being critical for generating the beneficial effects of prisms.
Consistent with the view that prisms influence the dorsal
stream, recent studies have shown dissociations in
neglect between benefits post prisms for motor tasks,
with unaltered performance on perceptual tasks [12,13].
Ferber and colleagues [13] used a chimeric faces (i.e., one
half smiling, the other neutral) task to show that before
prisms, fixations in a neglect patient were biased to the
right half of faces. When asked which of the two
chimerics seemed ‘happier’, the patient chose the face
smiling on the right – a bias opposite that of controls
[17]. After prisms the patient’s eye movements were now
biased towards fixating the left side of faces. Even though
the patient’s eye movements now explored the previously
neglected side of faces, he continued to choose the right-
smiling face as being ‘happier’. Although some suggest
that this may be specific to faces [18], others have shown
the same dissociation with simple shapes [12].
If prisms improve neglect by acting on the dorsal stream
there should be a greater effect on behaviors requiring
motor responses. In contrast, tasks relying on perception
436 Vision, central
0959-4965 c2010 Wolters Kluwer Health | Lippincott Williams & Wilkins DOI: 10.1097/WNR.0b013e328338592f
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
may be less likely to show benefits. We tested this using
the line bisection and landmark tasks – a perceptual
equivalent of bisection – in three neglect patients. The
landmark task requires patients to judge which end
of a horizontal line is closest to a bisection mark [19].
Responses reflect a perceptual judgment of the spatial
extent of the line. Patients consistently reporting that
the left end is closest to the bisection, even when it is at
center, reflects a perceptual distortion such that the left
half of the line is seen to be shorter. Although deficits on
bisection are considered to reflect the same bias, the fact
that this task requires a motor response distinguishes it
from the landmark task. That is, while performance on
both tasks reflects the same underlying perceptual bias,
improvement after prisms may only be expected for the
task that has a motor response (i.e., bisection).
Methods
Eight, right-handed, healthy, older individuals [three
males, mean age = 67 (± 5.5) years] with no history of
neurological or psychiatric illness were recruited from the
community. Three neglect patients were examined using
cancellation, figure copying, and bisection. For bisection,
deviations from center were calculated as a percentage of
total line length (leftward deviations coded as negative).
Neglect was considered to be present if mean bisections
deviated from center by 5% or more. For cancellation,
neglect was considered present if 10% or more of left
targets were omitted. Neglect was scored as present on
figure copying by visual inspection (Fig. 1).
NS was an 80-year-old, right-handed female who
presented with neglect after a stroke affecting right
parietal white matter and the right thalamus (Fig. 1). Her
bisections were biased rightward by 11% and had 75%
leftward omissions on cancellation. Neglect was evident
on figure copying. NS was right handed and had no
hemiplegia.
RR was a 66-year-old male who suffered a right hemi-
sphere stroke involving fronto-parietal white matter,
parietal cortex, and basal ganglia (Fig. 1). His bisection
was biased rightwards by 12%, had 90% leftward
omissions on cancellation, and neglect was evident on
figure copying. RR was right handed and was hemiplegic.
SQ was a 79-year-old female who suffered a stroke
affecting right occipito-temporal and medial parietal
cortex (Fig. 1). Her bisection was biased rightwards
by 12.8%, had 77% omissions of leftward targets on
cancellation, and neglect was evident on figure copying.
SQ was right handed, had a left hemianopia, and showed
no signs of hemiplegia.
All three patients showed a strong leftward bias on
the landmark task pre-PA (% of ‘left’ responses for
RR = 100%, NS = 100%, SQ = 90%) indicating they all
perceived the left half of the line to be shortest. Informed
consent was obtained from all patients before testing
and the protocol was approved by institutional ethics
committees in accordance with the Helsinki Declaration.
Apparatus and procedure
The bisection task consisted of 10 trials with a single
black line (length = 236 mm) presented horizontally on
a sheet of 8.5 1100 paper aligned to the patients’
midline. Patients used a pen to mark where they thought
center was.
The landmark task consisted of 16 black lines (length =
200 mm). Each line was placed horizontally on a sheet of
8.5 1100 paper. Ten lines had a vertical bisection mark
(height = 10 mm) placed at center, with six lines bisected
1, 3, or 5 mm to the left or right. Lines were presented
randomly and patients were told that no bisection
appeared at center. On half the trials, patients indicated
verbally which end of the line was closest to bisection.
On the other half of trials they indicated which end
was further away from bisection. This eliminated the
influence of response biases (for analysis, responses were
reverse coded when patients indicated which end was
further away). Only trials in which the bisection was at
center were included for analysis.
For PA, patients pointed to targets left and right of
midline while wearing prisms that shifted vision to 101
right. Patients pointed to targets every 2–3 s for 5 min.
Controls completed an identical procedure with left
shifting prisms as previous studies find robust effects
with this direction of shift in controls [20]. Controls were
adapted for 10 min as a previous study indicates that
longer exposure periods are needed for ‘neglect like’
patterns of performance to emerge in controls [20]. To
measure the effects of PA, patients and controls made
five pointing movements with eyes closed to a location
straight-ahead of their body (i.e., proprioceptive straight-
ahead judgment). These straight-ahead judgments were
made once before adaptation (baseline), once immedi-
ately postadaptation (post), and once at the end of
testing (late). Endpoints of movements were recorded by
the experimenter. For all tasks, patients and controls used
their right hand. Patients and controls first completed
one session of line bisection, landmark, and straight-
ahead pointing before being exposed to prisms, after
which they completed the same three tasks. Line
bisection and landmark tasks were counterbalanced
before and after PA. Straight-ahead pointing was repeated
a third time at the end of testing to determine whether
participants were still adapted.
Data analysis
Bisection was analyzed in terms of the percentage
deviation from center relative to total line length
(leftward deviations coded as negative). The landmark
Prisms influence action but not perception Striemer and Danckert 437
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
task was analyzed in terms of the percentage of ‘left’
responses (i.e., bisection was closer to the left end of
the line). For controls, to examine the effects of PA on
subjective straight-ahead, line bisection, and landmark
tasks, we ran paired samples t-tests. We did the same
thing for patients at both the group and individual
level. Finally, each patient’s performance pre and post-PA
was compared directly to controls using a modified
independent samples t-test, in which the patient is
treated as an individual sample not contributing to within
group variance [21]. Straight-ahead judgments were
measured as deviations from center (leftward deviations
coded as negative) and converted to degrees of visual
angle. Data were analyzed separately for each patient
using paired samples t-tests to contrast pre vs. post, and
pre vs. late straight-ahead pointing.
Fig. 1
Patient NS
Patient NS
Patient RR
Patient RR
Patient SQ
Patient SQ
Pre
Post
Late
Pre
Pre
Post
Post
Late
Late
−20246810
Degrees of visual angle
12 14
−20 2 4 6 810
Degrees of visual angle
12 14
−20 2 4 6 810
Degrees of visual angle
12 14
Patient scans (left), figure copying performance for the daisy (middle), and subjective straight-ahead judgments made preprisms (Pre), postprisms
(Post) and at the late session postprisms (Late). Patients NS (top) and RR (middle) both have computed tomography scans, whereas patient SQ
(bottom) has a sagittal MRI. Note that for both NS and RR scans are presented in radiological convention with the right hemisphere appearing on the
left side of the scan. Straight-ahead judgments are presented in degrees of visual angle (left is negative).
438 NeuroReport 2010, Vol 21 No 6
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
Results
Straight-ahead pointing
Controls
Straight-ahead judgments were shifted right after left
prisms [mean (standard error) pre = 3.4 (1.5) vs. mean
post = 10.3 (0.9), t(7) = 5.5, P= 0.001]. Controls had
begun to deadapt at the end of testing [post mean = 10.3
(0.9) vs. late mean = 6.9 (1), t(7) = 4.6, P< 0.01], how-
ever, straight-ahead judgments were still to the right of
judgments made pre-PA [pre mean = 3.4 (1.5) vs. late
mean = 6.9 (1), t(7) = 3.2, P< 0.05]. Thus, controls were
still adapted at the end of testing.
Straight-ahead judgments for two of the three patients
(RR and SQ) were shifted leftward immediately post-PA,
and all three patients were shifted significantly left-
ward at the late test (all P< .05; Fig. 1). Thus, at the
last test all patients were still significantly adapted to
prisms.
Line bisection
For controls pre-PA bisections were slightly left of center
(mean bias = – 0.21%). In all but one control, mean
bisections were shifted right post-PA [mean bias = 1.43%,
t(7) = 3.5, P< 0.01; Fig. 2].
For all three patients there was a leftward shift in
bisection post-PA [NS, t(9) = 2.8, P<0.05; RR,t(9) = 3.2,
P< 0.05; SQ, t(9) = 8.6, P< 0.001; Fig. 2]. In addition,
Crawford’s t-test on pre-PA bisections indicated that all
three patients were well outside the range of controls
(Table 1). When postexposure performance was com-
pared in the same way, results showed that all three
patients were well within the range of controls (Table 1).
Landmark task
For controls the percentage of ‘left’ responses was 63.8%
pre-PA (Fig. 2). This was not significantly altered post-PA
at the group level [mean % (standard error) ‘left’
responses =71.25% (10.3), t(7) = – 1.2, P=0.3]. To ex-
amine whether the null effect on landmark performance
resulted from deadaptation before completion of the
bisection task (owing to the motor response required), we
compared landmark performance pre vs. post-PA for
controls who completed the bisection task first post-PA,
or the landmark task first post-PA. This analysis revealed
a significant increase in ‘left’ responses post-PA for
controls who completed the landmark task first post-PA
[pre–post difference = 18%; t(4) = 3.09, P= 0.037] com-
pared with a tendency to make fewer ‘left’ responses
in those who completed the landmark task after bisec-
tion post-PA [pre–post difference = – 10%; t(2) = 1.73,
P= 0.23; Fig. 2].
Two patients (NS and RR) responded ‘left’ on 100% of
trials pre and post-PA. SQ responded ‘left’ on 87.5% of
trials pre-PA and on 90% of trials post-PA. Thus, all three
patients showed a strong bias towards seeing the left half
of lines as shorter than the right, a bias that was unaltered
by PA (Fig. 2). Note that deadaptation cannot explain the
null effect in patients as adaptation remained stable,
or significantly increased by the late test in all three
patients (Fig. 1).
Discussion
We examined the effect of PA on perceptual and motor
performance using the bisection and landmark tasks in
three neglect patients and a group of controls. Results
showed that for neglect patients, PA reduced rightward
biases in bisection, but had no effect on leftward
perceptual biases on the landmark task (Fig. 2). Thus,
despite the fact that bisections were shifted in the
opposite direction to the prism shift, there was no
influence on how the spatial extent of lines was per-
ceived. The dissociation between perceptual and motor
influences of PA shown here is consistent with other
studies showing that PA shifts eye movements left
without altering the perception of stimuli in neglected
space [12,13]. These data are consistent with the notion
that PA influences dorsal stream processes without
influencing perceptual biases in the ventral stream in
patients with neglect.
Performance on line bisection and landmark tasks in
younger healthy controls typically reveal a leftward over-
estimation of line length (i.e., pseudoneglect) that results
in a leftward bias in bisection, and a tendency to say the
prebisection mark is closer to the right end of the line.
Consistent with earlier studies on perceptual biases in
elderly patients [22,23], our controls showed only a small
leftward bias in bisection and no bias in the landmark
task. Irrespective of the presence of pseudoneglect
before PA, leftward PA resulted in a ‘neglect-like’ pattern
of performance for line bisection (i.e., a rightward shift)
and, for those who completed the landmark task first
postprisms, an increase in the percentage of ‘left’
responses in the landmark task. This is consistent with
earlier studies indicating that prisms can influence both
perceptual and motor biases in healthy individuals
[20,24,25]. These results are informative because they
suggest that the landmark task used here was sensitive
enough to detect any effects of PA in patients where such
effects existed. In addition, they suggest it is possible
for prisms to influence perception in individuals without
brain damage, perhaps through intact connections be-
tween the dorsal and ventral streams. Importantly, the
failure to alter perceptual biases in the face of changes
to visuomotor performance in neglect may be precisely
because of the fact that damage to inferior parietal cortex
disrupts the normal interaction between the dorsal and
ventral streams [6], eliminating the path by which prisms
could influence perception in neglect.
Prisms influence action but not perception Striemer and Danckert 439
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
Fig. 2
Healthy controls
Line bisection
Pre
Post
Pre
Pre Pre
Post
Post
−1−0.5 0.5 1.5 2.5 100 80 60 40 20 0321
1
8
6
4
2
0
−2
−4
100
80
60
40
20
0
100
80
60
40
20
0
−2
100
80
60
40
20
0
Percentage of ‘left’ responses
Deviation (percentage of line length)
Deviation (percentage of line length)
Percentage of ‘left’ responses
2345678
−20 2 4 6 8 10 12 14 16 18 100 80 60 40 20 0
Deviation (percentage of line) Percentage of ‘left’ responses
Percentage of ‘left’ responses
Participants
Deviation (percentage of line length)
Participants
Line bisection
SQ NS RR
Participants
SQ NS RR
Neglect patients
12345678
Participants
0
NS
NS
∗
∗
∗
∗
∗
∗
∗
∗∗
∗
Landmark
Landmark
Post
Pre Post
Pre Post
Upper panel: healthy older control data. Mean [standard error (SE)] line bisection performance pre (open circle) and postadaptation (filled circle)is
to the left, with mean (SE) percentage of ‘left’ responses for the landmark task pre (open bar) and post (filled bar) adaptation to the right. Below are
the data for individual participants on each task. *The participant performed the landmark task first post adaptation. Lower panel: neglect patient
data: mean (SE) line bisection performance (i.e., deviation from center represented as a percentage of total line length, rightward deviations codedas
positive) pre (open circles) and postprisms (filled black circles) is presented on the left. Mean (SE) percentage of ‘left’ responses for pre (open bar)
and postprisms (filled bar) is presented to the right. Below this are the individual performances of patients with neglect on each task. *Significant
difference at P< 0.05.
440 NeuroReport 2010, Vol 21 No 6
Copyright © Lippincott Williams & Wilkins. Unauthorized reproduction of this article is prohibited.
Conclusion
Many of the beneficial effects of prisms observed in
neglect may arise through interactions with the dorsal
‘action’ stream but may fail to influence perceptual biases
evident in neglect.
Acknowledgements
This study was supported by Heart and Stroke Founda-
tion of Canada Postdoctoral award, and a Natural Sciences
and Engineering Research Council of Canada PhD award
to C.S. This study was also supported by a Heart and
Stroke Foundation of Canada Grant-in-Aid and Natural
Sciences and Engineering Research Council of Canada
Discovery and Canada Research Chair (Tier II) awards
to J.D. The authors would like to thank Phil Hatcher for
his assistance with collecting the control data.
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Table 1 Mean (standard error) line bisection performance
(deviation from centre as a % of total line length; rightward
deviations codes as positive) for controls and each patient
pre and postprisms
Preprisms Postprisms
Control, mean (SD) – 0.21 (3.05) 1.43 (3.2)
Patient NS 11.02 0.88
Crawford’s t-test t= – 10.496, P= 0.005 t=0.49, P=0.438
Pbelow 99.49% 43.79%
Patient RR 6.81 – 0.98
Crawford’s t-test t=–6.56,P= 0.033 t= 2.13, P= 0.25
Pbelow 96.68% 25.07%
Patient SQ 18.98 3.49
Crawford’s t-test t=–17.54,P= 0.0003 t= – 1.82, P= 0.28
Pbelow 99.97% 71.72%
‘Pbelow’ refers to the percentage of the healthy population with scores falling
below the patient’s as determined by Crawford’s t-test.
SD, standard deviation.
Prisms influence action but not perception Striemer and Danckert 441
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