Neuropsychologia 141 (2020) 107314

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Neuropsychologia
journal homepage: http://www.elsevier.com/locate/neuropsychologia

Eye dominance modulates visuospatial attention

S. Schintu a, b, c, d, *, R. Chaumillon e, A. Guillaume f, R. Salemme a, b, g, K.T. Reilly a, b, L. Pisella a, b,
A. Farn�e a, b, g, h
a
Integrative Multisensory Perception Action & Cognition Team (ImpAct), INSERM U1028, CNRS UMR5292, Lyon Neuroscience Research Center (CRNL), Lyon, France
b
University of Lyon 1, Lyon, France
c
Department of Psychology, George Washington University, WA, USA
d
Behavioral Neurology Unit, National Institute of Neurological Disorders and Stroke, National Institutes of Health, Bethesda, MD, USA
e
Aix Marseille Universit�e, CNRS, LNC UMR 7291, FR 3C FR 3512, Marseille, France
f
Paris Descartes University, Vision Action Cognition Laboratory, EA7326, Institute of Psychology, Boulogne-Billancourt, Paris, France
g
Hospices Civils de Lyon, Mouvement & Handicap, Neuro-Immersion Platforms, Lyon, France
h
Center for Mind/Brain Sciences (CIMeC), University of Trento, Italy

A R T I C L E I N F O A B S T R A C T

Keywords: Visuospatial attention has an inherent asymmetry: the leftward bias called pseudoneglect. In typical line
Pseudoneglect bisection tasks, healthy individuals tend to judge the center of a line leftward of the true center, an effect
Asymmetry attributed to the right hemisphere dominance in visuospatial attention. Since it has been shown that information
Laterality
perceived by the dominant eye strongly activates the ipsilateral visual cortex, we hypothesized that eye domi­
Visual cortex
Line bisection
nance may modulate visuospatial attention bias. Because activation of the left hemisphere induced by left eye
dominance should mitigate the right hemisphere dominance in attention, we predicted that right-handed in­
dividuals with left dominant eye would show smaller amount of pseudoneglect than right-handed individuals
with right dominant eye. We compared the performance at both the perceptual (Landmark) and manual line
bisection task of forty right-handed healthy individuals, half of whom had a right dominant eye and the other
half a left dominant eye. As predicted, the left eyed dominant group showed smaller, actually not significant
pseudoneglect, which was thus greater in the right eye dominant group. The influence of eye dominance on
visuospatial attention was present in the Landmark but not the manual line bisection task, in which the amount
of visuospatial bias correlated with participants’ degree of (right) handedness. This is the first report of the effect
of eye dominance on visuospatial attention within a right-handed population. This finding, by showing the in­
fluence of eye dominance on visuospatial cognition, not only helps in better defining intact visuospatial cognition
mechanism but also encourages further research to pinpoint the neural basis of such interaction.

1. Introduction as the activation-orientation theory posits, this increased attention to

the contralateral (left) hemifield makes the left visual space (i.e., left
Young individuals free of neurological disorders usually judge the side of the line) being perceived longer then the right one, and thus
center of a line leftward of the true center as measured by line bisection judging the center of line leftward of the true center (Bultitude and
tasks (Milner et al., 1992; Schenkenberg et al., 1980). This leftward bias Aimola Davies, 2006).
in intact visuospatial attention is called pseudoneglect (Bowers and Pseudoneglect is typically present at the population level, but much
Heilman, 1980; for a review see Jewell and McCourt, 2000). Pseudo­ inter individual variability is observed (Manning et al., 1990; McCourt
neglect is hypothesized to be the product of the uneven distribution of and Olafson, 1997; Jewell and McCourt, 2000; Toba et al., 2011;
attention due to the right hemispheric dominance in attention (Kins­ Szczepanski and Kastner, 2013). Among the several factors influencing
bourne, 1970; Mesulam, 1999; Nicholls and Roberts, 2002). The right spatial attention performance, such as handedness strength (Ochando
hemisphere is indeed the most activated one during attentional task and, and Zago, 2018), eye dominance is gaining increasing interest (e.g.,

* Corresponding author. Behavioral Neurology Unit, National Institute of Neurological Disorders and Stroke, National Institutes of Health, 10 Center Drive,
Bethesda, MD, USA.
E-mail addresses: [email protected], [email protected] (S. Schintu).

https://doi.org/10.1016/j.neuropsychologia.2019.107314
Received 21 July 2019; Received in revised form 16 December 2019; Accepted 18 December 2019
Available online 21 December 2019
0028-3932/Published by Elsevier Ltd.
S. Schintu et al. Neuropsychologia 141 (2020) 107314

Roth et al., 2002; Szczepanski and Kastner, 2013). Eye dominance and dominance to manifest its influence on visuospatial bias when assessed
handedness have been shown to have a very weak relationship, with in the more perceptual manner, i.e., the Landmark task, and handedness
66% of right-handers having a right DE and 60% of left-handers having a to show its influence on visuospatial bias when assessed by manual line
left DE (Bourassa et al., 1996). Numerous studies failed to show a clear bisection because of the motor bias intrinsic to the task.
link between the two, which is not surprising given the fact the affer­ Our model and predictions were supported by the results, showing
ence/efference of one hand is associate to one hemisphere whereas the that eye dominance contributes in determining the presence and amount
efference of one eye is associated to both hemispheres (Mapp et al., of pseudoneglect in the precited direction solely when assessed by the
2003). The dominant eye is defined as the eye manifesting physiological Landmark task. This finding may help account for the inter-individual
superiority and it is the one chosen to align a target in peripersonal space variability observed in the pseudoneglect phenomenon.
with a more distant point, or when performing a monocular task such as
looking through a small hole (it is referred to as sighting dominant eye; 2. Materials and methods
Porac and Coren, 1976). There are interocular differences in activation
of visual areas in response to stimuli presented to the contralateral visual 2.1. Participants
field, with the dominant eye generating a stronger activation in response
to sensory information as compared to the one generated by the Forty adults with normal or corrected-to-normal vision and no his­
non-dominant one (Mendola and Conner, 2007; Rombouts et al., 1996). tory of neurological problems were recruited for the study. All partici­
Since the dominant eye, as compared to the non-dominant one, activates pants gave informed consent and were right-handed according to the
larger portions of the ipsilateral primary visual cortex (Erdogan et al., Edinburgh Handedness Inventory, which is used to quantified the degree
2002), it has been concluded that ocular dominance is mainly depending of handedness (Oldfield, 1971). The study was approved by the French
on the ipsilateral occipital lobe (Shima et al., 2010) possibly as a result of national ethics committee (CPP SUD EST IV) and was conducted in
more numerous, or stronger inputs gathered by the temporal hemiretina accordance with the ethical standards of the 1964 Declaration of Hel­
and projecting to the ipsilateral visual cortex. Consistent with this last sinki. Twenty participants were in the right dominant eye (RDE) group
proposition, simple reaction times in response to a lateralized visual and the other twenty in the left dominant eye (LDE) group. In order to
target are shorter for the contralateral visual hemifield with respect to have comparable numbers in the two groups, participants were screened
the DE (Chaumillon et al., 2014). Globally, there is an advantage for for eye dominance by having them perform three repetitions of the
stimuli processed by the dominant eye: perceived image size is increased hole-in-card test (Miles, 1930). The number of participants for each
(Coren and Porac, 1976) and percept is more salient, suggesting that group was set to 20; once one group was complete recruitment
information from the dominant eye has priority in visual processing continued for the other group. All participants had unambiguous eye
(Shneor and Hochstein, 2008, 2006). dominance, as for all forty participants the hole in the card was aligned
Grounding on the activation-orientation theory (Reuter-Lorenz et al., with the same eye on each repetition. Two participants were excluded
1990) we hypothesized that eye dominance may impact physiologically because the Edinburgh Handedness Inventory score was missing. The
visuospatial attention bias, and namely that individuals with right final sample submitted to statistical analysis was composed by
dominant eye should show a greater amount of pseudoneglect as thirty-eight participants: eighteen in the RDE group (4 males, mean age
compared to individuals with a left dominant eye. The activation- ¼ 26.4, SEM ¼ 0.99, SD ¼ 4.20) and the remaining twenty in the LDE
orientation theory, by postulating that the distribution of attention is group (9 males, mean age ¼ 28.4 SEM ¼ 1.18, SD ¼ 5.25). The two
biased toward the direction opposite to the most activated hemisphere, groups did not differ in terms of age [t(36) ¼ 1.257 p ¼ 0.217], and
accounts for the pseudoneglect phenomenon as due to a biased similarly the degree of Handedness did not differ between the RDE
perception of the half of the line in the left hemifield because of the right (mean ¼ 87.25, SEM ¼ 3.46, SD ¼ 14.70) and LDE (mean ¼ 80.06, SEM
hemisphere dominance for visuospatial attention. We predicted that ¼ 4.30, SD ¼ 19.23) group [t(36) ¼ 1.282 p ¼ 0.208].
right-handed individuals with right dominant eye, as compared to right- Following the eye dominance test, participants performed both a
handed individuals with left dominant eye, should show a greater perceptual (Landmark task; Milner et al., 1992) and manual (Schen­
amount of pseudoneglect. This would depend on of the fact that infor­ kenberg et al., 1980) line bisection task, the order of which was coun­
mation from the left side of the line is perceived elongated because it is terbalanced across participants. During both tasks, which were carried
processed by the right hemisphere, which not only is the most activated out in binocular vision, participants were seated comfortably with their
by the attentional task, but also by the inputs of their right dominant eye head positioned on a chinrest.
to the right visual cortex (i.e., right hemisphere dominance plus domi­
nant eye). By contrast, in individuals with left dominant eye, the left­ 2.2. Landmark task
ward bias would be mitigated by the activation of the left hemisphere
due to stronger inputs from the left dominant eye to the left visual A modified version of the Landmark task (Milner et al., 1992) was
cortex. We thus put forward here a simple model to predict the mani­ used to quantify visuospatial performance. The Landmark task (as in
festation of pseudoneglect as a function of eye dominance: left eye Schintu et al., 2014) consisted of a series of pre-bisected lines presented
dominance, by increasing the activity in the left visual cortex, would centrally on a computer screen (22 inches (559 mm � 274 mm), res­
dampen the typical bias depending upon right hemisphere dominance. olution 1680 � 1050, refresh rate 60 HZ) positioned with the center
We tested this prediction by comparing the performance at both the aligned to the participant’s mid-sagittal axis. Participants placed their
perceptual (Landmark) and manual line bisection tasks, between one hands on their lap beneath the table and were instructed to fully inspect
group of healthy right-handed participants with right dominant eye and each pre-bisected line and judge whether the mark (vertical transector)
one group of healthy right-handed participants with left dominant eye. was closer to the left or right end of the line. In this two-alternative
These tasks, despite being both used to quantify visuospatial asymmetry forced-choice paradigm participants answered by pressing either the
(Zago et al., 2017), bear substantial differences (Milner et al., 1992). The left (transector closer to the left end of the line) or right (transector
Landmark task can be summarized as a pure test of perceptual/atten­ closer to the right end of the line) pedal positioned under each foot. They
tional bias, as it requires to judge a series of pre-bisected lines and, in our were instructed to respond as accurately and quickly as possible. Prior to
study, avoids any hand movement, whereas the motor line bisection task the task, at least ten practice trials were given to ensure that participants
involves motor bias by requiring to actively bisect a line with the correctly understood the instructions and were confident answering
dominant hand. We therefore expect eye dominance and strength of with the pedals. The stimuli were white lines (350 mm x ~2 mm) dis­
handedness to have a different weight in the quantification of the vi­ played on a black screen positioned 35 cm from the participant’s eyes.
suospatial bias according to the task used to measure it. We predict eye Lines were transected at the true center and at 2, 4, 6, 8, and 10 mm

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S. Schintu et al. Neuropsychologia 141 (2020) 107314

toward the left and right of the true center (0 mm). Each of the 11 not [F(1,35) ¼ 0.021, p ¼ 0.886, η2p ¼ 0.001].
different pre-bisected lines was presented six times in a random order, The subsequent independent univariate ANOVAs we ran to follow up
yielding a total of 66 trials. Each pre-bisected line was displayed for a the Task � Group interaction revealed that, as quantified by the Land­
maximum of 5 s or until a response was made and was then replaced by a mark task, the RDE group had a greater amount of pseudoneglect (mean
black-and-white patterned mask which stayed on the screen for 1 s ¼ 1.85 mm, SEM ¼ 0.46) as compared to the LDE group (mean ¼
before the next pre-bisected line was displayed. Presentation software 0.28 SEM ¼ 0.34) [F (1, 36) ¼ 6.083 p ¼ 0.019, η2p ¼ 0.145] (Fig. 1).
(Neurobehavioral Systems, Inc., USA) was used to generate the stimuli, In contrast, no difference was found between the RDE (mean ¼ 0.81
record responses, and control the timing of stimulus presentation SEM ¼ 0.87) and LDE (mean ¼ 0.12 SEM ¼ 0.7) groups at the manual
throughout the task. For each participant, the percentage of ‘right’ re­ line bisection task [F (1, 36) ¼ 0.423 p ¼ 0.520] (Fig. 1). Moreover, the
sponses was plotted as a function of the position of the transector. These amount of pseudoneglect as quantified by the Landmark task was
data were then fitted with a sigmoid function and the value on the x-axis significantly different from zero for the RDE group [t(17) ¼ 4.028 p ¼
corresponding to the point at which the participant responded ‘right’ 0.001], but not for the LDE group [t(19) ¼ 0.635 p ¼ 0.533] con­
50% of the time was taken as that participant’s point of subjective firming the presence of pseudoneglect solely for the RDE group.
equality (PSE). To follow up the Task x Degree of Handedness interaction we
This task prioritizes the perceptual and minimizes the motor computed two Spearman correlations between the degree of handedness
component of the visuospatial bias by asking participants to judge a and the performance at each task. These analyses revealed a significant
series or pre-bisected lines instead of actively bisect them (Milner et al., correlation between the degree of handedness and the performance
1992). solely for the Manual line bisection (r ¼ 0.516 p ¼ 0.001), and not for
the Landmark task (r ¼ 0.012 p ¼ 0.941) (Fig. 2). Based on (Ochando
2.3. Manual line bisection task and Zago, 2018) finding, we categorized participants based on their
degree of Handedness, independently of their eye dominance. Partici­
The manual line bisection task (Schenkenberg et al., 1980), consisted pants whose score at the Edinburgh task was >85 were categorize as
of a series of 10 black lines (350 mm x ~2 mm which were of identical in “strong” righthanded (n ¼ 21) and those whose score was <85 as “weak”
size to those used for the Landmark task) each drawn on A3 (297 mm � right-handed (n ¼ 17). The independent t-test comparing the motor line
420 mm) sheets of paper that were positioned over the computer screen bisection performance revealed that the strong right-handed group
which was kept at the same distance and position as for the Landmark showed a leftward bias (mean ¼ 1.23 SD ¼ 4.20 SEM ¼ 0.92) and the
task. Participants were instructed to fully inspect each line and with the weaker right-handed group showed a rightward bias (mean ¼ 2.24 SD ¼
pen held in their right hand, draw a vertical mark where they thought 3.78 SEM ¼ 0.92) [t(36) ¼ 2.645 p ¼ 0.012, Cohens d ¼ 0.87],
the center of the line to be. Once the mark had been drawn the experi­ whereas no difference was found between the strong and weak hand­
menter then turned the page to reveal the following line. No time limit edness groups’ performance at the Landmark task [t(36) ¼ 0.475 p ¼
was imposed, and participants took on average 1 s to place the mark on 0.638].
each line. For each of the ten lines the distance between the mark placed These results show that in right handers, eye dominance did not
by the participant and the true center of the line was calculated. The PSE affect performance on a manual line bisection task. Indeed, participants
was calculated as the average across the ten trials of the distance be­ in both the RDE and LDE groups were very accurate when marking the
tween the true center and the mark, with marks to the right of center center of the line with their right hand. In contrast, on the Landmark task
coded as positive and to the left as negative. the performance of the two groups differed significantly; the LDE group
This task, while perceptual in nature, by asking participants to accurately judged the center close to the true center, whereas the RDE
actively bisect a series of lines may nullify the perceptual bias because of group exhibited a significant leftward bias as they judged the center to
the kind motor response involved (Milner et al., 1992). be significantly leftward of the true center.

2.4. Statistical analyses 4. Discussion

Statistical analyses were carried out using the SPSS software package We hypothesized that eye dominance impacts visuospatial attention,
(Version 24, Chicago, IL: SPSS Inc.). The significance threshold was set and thus that the asymmetric hemisphere activation over the visual
at 0.05 (two-tailed). All values in the text are presented with standard areas triggered by eye dominance interacts with the typical right
error of the mean (SEM) and when appropriate also standard deviation hemisphere activation in attentional task. The current findings support
(SD) and effect size are reported. The performances of the Landmark and the hypothesis that eye dominance influences visuospatial attention
Manual line bisection tasks were submitted to repeated measure Anal­ mechanisms and, as predicted, shows that right-handed individuals with
ysis of Variance (ANOVA) having Task (Manual line bisection, Land­ right dominant eye exhibit a greater amount of pseudoneglect compared
mark) as within variable, Group (RDE, LDE) as between variable and to individuals with left eye dominance.
Degree of Handedness (i.e., Edinburgh score) as covariate. We included The right dominant eye group showed a significant leftward bias and
degree of handedness as covariate because it has been shown to a greater amount of pseudoneglect than the group with left dominant
modulate line bisection performance (Jewell and McCourt, 2000). eye only when measured by the Landmark task, but not by the manual
line bisection task. The Landmark task (Milner et al., 1992) is typically
3. Results administered to healthy participants whereas the manual line bisection
(Schenkenberg et al., 1980) is the most common bedside tool for hem­
The repeated measure ANOVA revealed a main effect of Task [F ispatial neglect screening. These two tasks are implied to measure the
(1,35) ¼ 21.640, p < 0.001, η2p ¼ 0.382] such that the PSE was more same phenomenon and are anecdotally compared across populations,
leftward (i.e., negative) in the Landmark task (mean ¼ 1.02 mm; SEM ¼ despite the effective correlation between them has never been produced
0.34) than in the Manual line bisection task (mean ¼ 0.32 mm; SEM ¼ and differences between the two have been reported in the literature.
0.70). The Task � Group interaction [F(1,35) ¼ 9.030, p ¼ 0.005, η2p ¼ The difference in performance at the two tasks can lie in the amount of
0.205] was significant, meaning that after we controlled for Degree of variability present in the manual line bisection performance and its poor
Handedness there was a significant difference in the performance be­ sensitivity (McCourt and Jewell, 1999; McCourt and Olafson, 1997) as
tween the RDE and LDE groups according to the task performed. The compared to the Landmark task. The Landmark task is thought to
Task x Degree of Handedness interaction was also significant [F(1,35) ¼ minimize the motor component and be more sensitive to the perceptual
18.084, p < 0.001, η2p ¼ 0.341] whereas the main effect of group was one (Milner et al., 1992), providing finer quantification of visuospatial

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S. Schintu et al. Neuropsychologia 141 (2020) 107314

Fig. 1. Point of Subjective Equality (PSE) at the Landmark and Manual line bisection tasks as a function of eye dominance. Negative values represent a PSE to the left
of the true center (pseudoneglect). Values are means and error bars represent the SEM.

Fig. 2. Pearson correlation between Edinburgh scale score and manual line bisection task (panel A) and Landmark task (panel B).

attention bias than manual line bisection (McIntosh et al., 2019). The the veridical center (Bultitude and Aimola Davies, 2006). However, the
greater amount of pseudoneglect observed with the Landmark task is the biased perception of stimuli in the left hemifield, due to the increased
consequence of the elimination, minimization, and control of con­ activation of the right hemisphere in the attentional task, interacts with
founding factors such as the motor responses associated with the manual the priority in visual processing for stimuli processed by the dominant
version of the line bisection (McCourt and Jewell, 1999; McCourt and eye (Shneor and Hochstein, 2008, 2006) that is the result of the
Olafson, 1997). Our results, by showing that the influence of eye increased activation of the visual cortex ipsilateral the to the dominant
dominance was measurable only via the Landmark task (devoid of any eye (Shima et al., 2010). In participants with right dominant eye, the
hand movement) are coherent with such argument. The additional special relationship between the dominant eye and its ipsilateral hemi­
finding that the correlation between the degree of handedness and the sphere (i.e., the right hemisphere; Shima et al., 2010) is consistent with
amount of pseudoneglect was significant for the manual line bisection the right asymmetry of the attentional networks. In contrast, in partic­
tasks, but not for the Landmark task, provides further support to this ipants with left dominant eye, the privileged relationship between the
conclusion. Conclusion that is even further strengthened by the fact that, dominant eye and its ipsilateral hemisphere (i.e., the left hemisphere)
when participants were categorized and compared by handedness counteracts the right hemisphere dominance for visuospatial attention,
strength, difference in performance was observed for the manual line possibly resulting in a smaller amount of pseudoneglect.
bisection and not for the Landmark task. The advocated involvement of primary visual areas in higher
Our model of eye-dominance dependent modulation of attentional cognitive functions is not surprising since the traditional interpretation
biases was grounded on the activation–orientation theory (Reuter-Lor­ of the activity in the primary visual cortex has been challenged by
enz et al., 1990). This theory by postulating that the distribution of findings showing that reward-timing activity can occur very early in
attention is biased to the direction opposite to the more activated sensory-processing paths such in the dominant eye visual area (Shuler
hemisphere, offers an account for the pseudoneglect origin. Since vi­ and Bear, 2006). When considering the possible neural underpinning of
suospatial tasks such as line bisection involve activation of the right the behavioral finding reported here, it is crucial to consider the key role
hemisphere (Fink et al., 2000) the contralateral (left) half of the line is of posterior parietal cortex (PPC) in the frontoparietal network, since its
the focus of greater attention and perceived as longer relative to the less modulation by transcranial direct-current stimulation directly impacts
attended right half, resulting in midpoint judgment located to the left of the amount of pseudoneglect (i.e., in a Landmark task, Ribolsi et al.,

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S. Schintu et al. Neuropsychologia 141 (2020) 107314

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