Gaze data of 4243 participants shows link between leftward and superior attention biases and age.

Healthy individuals typically show more attention to the left than to the right (known as pseudoneglect), and to the upper than to the lower visual field (known as altitudinal pseudoneglect). These biases are thought to reflect asymmetries in neural processes. Attention biases have been used to investigate how these neural asymmetries change with age. However, inconsistent results have been reported regarding the presence and direction of age-related effects on horizontal and vertical attention biases. The observed inconsistencies may be due to insensitive measures and small sample sizes, that usually only feature extreme age groups. We investigated whether spatial attention biases, as indexed by gaze position during free viewing of a single image, are influenced by age. We analysed free-viewing data from 4,243 participants aged 5-65 years and found that attention biases shifted to the right and superior directions with increasing age. These findings are consistent with the idea of developing cerebral asymmetries with age and support the hypothesis of the origin of the leftward bias. Age modulations were found only for the first seven fixations, corresponding to the time window in which an absolute leftward bias in free viewing was previously observed. We interpret this as evidence that the horizontal and vertical attention biases are primarily present when orienting attention to a novel stimulus - and that age modulations of attention orienting are not global modulations of spatial attention. Taken together, our results suggest that attention orienting may be modulated by age and that cortical asymmetries may change with age.

Top-down control of the left visual field bias in cued visual spatial attention.

A left visual field (LVF) bias in perceptual judgments, response speed, and discrimination accuracy has been reported in humans. Cognitive factors, such as visual spatial attention, are known to modulate or even eliminate this bias. We investigated this problem by recording pupillometry together with functional magnetic resonance imaging (fMRI) in a cued visual spatial attention task. We observed that (i) the pupil was significantly more dilated following attend-right than attend-left cues, (ii) the task performance (e.g. reaction time [RT]) did not differ between attend-left and attend-right trials, and (iii) the difference in cue-related pupil dilation between attend-left and attend-right trials was inversely related to the corresponding difference in RT. Neuroscientically, correlating the difference in cue-related pupil dilation with the corresponding cue-related fMRI difference yielded activations primarily in the right hemisphere, including the right intraparietal sulcus and the right ventrolateral prefrontal cortex. These results suggest that (i) there is an asymmetry in visual spatial attention control, with the rightward attention control being more effortful than the leftward attention control, (ii) this asymmetry underlies the reduction or the elimination of the LVF bias, and (iii) the components of the attentional control networks in the right hemisphere are likely part of the neural substrate of the observed asymmetry in attentional control.

Pseudoneglect during object search in naturalistic scenes.

Pseudoneglect, that is the tendency to pay more attention to the left side of space, is typically assessed with paper-and-pencil tasks, particularly line bisection. In the present study, we used an everyday task with more complex stimuli. Subjects' task was to look for pre-specified objects in images of real-world scenes. In half of the scenes, the search object was located on the left side of the image (L-target); in the other half of the scenes, the target was on the right side (R-target). To control for left-right differences in the composition of the scenes, half of the scenes were mirrored horizontally. Eye-movement recordings were used to track the course of pseudoneglect on a millisecond timescale. Subjects' initial eye movements were biased to the left of the scene, but less so for R-targets than for L-targets, indicating that pseudoneglect was modulated by task demands and scene guidance. We further analyzed how horizontal gaze positions changed over time. When the data for L- and R-targets were pooled, the leftward bias lasted, on average, until the first second of the search process came to an end. Even for right-side targets, the gaze data showed an early left-bias, which was compensated by adjustments in the direction and amplitude of later saccades. Importantly, we found that pseudoneglect affected search efficiency by leading to less efficient scan paths and consequently longer search times for R-targets compared with L-targets. It may therefore be prudent to take spatial asymmetries into account when studying visual search in scenes.

Visuospatial perception is not affected by self-related information.

Previous research suggests that attention is drawn by self-related information. Three online experiments were conducted to investigate whether self-related stimuli alter visuospatial perceptual judgments. In a matching task, associations were learned between labels ('Yourself'/friend/stranger's name) paired with cues. Cues were coloured outlines (Experiment 1, N = 135), geometric shapes (Experiment 2, N = 102), or coloured gradients (Experiment 3, N = 110). Visuospatial perception bias was measured with a greyscales task. Cues were presented prior to, and/or alongside greyscales. We hypothesized there would be a bias towards the self-related cue. In all experiments, we found a self-related bias in the matching task. Furthermore, there was an overall leftward visuospatial perceptual bias (pseudoneglect). However, we found anecdotal to moderate evidence for the absence of an effect of self-related cues on visuospatial perception judgments. Although self-related stimuli influence how our attention is oriented to stimuli, attention mechanisms that influence perceptual judgements are seemingly not affected by a self-bias.

Smartphones and rightward collisions.

People tend to deviate to the right when walking through a narrow aperture (e.g., a doorway), resulting in a rightward bias in collisions. This study examines the effects of smartphone use on rightward collisions while walking. When pedestrians walk through a narrow aperture, they usually head straight to the perceived centre of the aperture, which is shifted slightly to the right, without updating the estimates. The rightward shift of the perceived centre is attributable to the rightward attentional shift in the extrapersonal space. Pedestrians using smartphones tend to fixate on the phone most of the time and thus tend not to look at their surroundings (i.e., extrapersonal space). Therefore, we predict that smartphone use will reduce rightward collisions. To test this prediction, we used a narrow-doorway task in which participants walked through a narrow doorway either with or without a smartphone. The participants with smartphones used them to perform either verbal or spatial tasks. The number of rightward collisions decreased when the participants used smartphones. The type of task had no effect on the lateral collision biases. These results were interpreted in terms of lateral attentional bias in peripersonal and extrapersonal spaces.

A meta-analysis of the line bisection task in children.

Meta-analyses have shown subtle, group-level asymmetries of spatial attention in adults favouring the left hemispace (pseudoneglect). However, no meta-analysis has synthesized data on children. We performed a random-effects meta-analysis of spatial biases in children aged ≤16 years. Databases (PsycINFO, Web of Science & Scopus) and pre-print servers (bioRxiv, medRxiv & PsyArXiv) were searched for studies involving typically developing children with a mean age of ≤16, who were tested using line bisection. Thirty-three datasets, from 31 studies, involving 2101 children, were included. No bias was identified overall, but there was a small leftward bias in a subgroup where all children were aged ≤16. Moderator analysis found symmetrical neglect, with right-handed actions resulting in right-biased bisections, and left-handed actions in left-biased bisections. Bisections were more leftward in studies with a higher percentage of boys relative to girls. Mean age, hand preference, and control group status did not moderate biases, and there was no difference between children aged ≤7 and ≥7 years, although the number of studies in each moderator analysis was small. There was no evidence of small study bias. We conclude that pseudoneglect may be present in children but is dependent on individual characteristics (sex) and/or task demands (hand used).Registration: Open Science Framework (https://osf.io/n68fz/).

Accuracy and precision of responses to visual analog scales: Inter- and intra-individual variability.

Visual analog scales (VASs) are gaining popularity for collecting responses in computer administration of psychometric tests and surveys. The VAS format consists of a line marked at its endpoints with the minimum and maximum positions that it covers for respondents to place a mark at their selected location. Creating the line with intermediate marks along its length was discouraged, but no empirical evidence has ever been produced to show that their absence does any good. We report a study that asked respondents to place marks at pre-selected locations on a 100-unit VAS line, first when it only had numerical labels (0 and 100) at its endpoints and then when intermediate locations (from 0 to 100 in steps of 20) were also labeled. The results show that settings are more accurate and more precise when the VAS line has intermediate tick marks: The average absolute error decreased from 3.02 units without intermediate marks to 0.82 units with them. Provision of intermediate tick marks also reduced substantially inter- and intra-individual variability in accuracy and precision: The standard deviation of absolute error decreased from 0.87 units without tick marks to 0.25 units with them and the standard deviation of signed distance to target decreased from 1.16 units without tick marks to 0.24 units with them. These results prompt the recommendation that the design of VASs includes intermediate tick marks along the length of the line.

A Meta-Analysis of Line Bisection and Landmark Task Performance in Older Adults.

Young adults exhibit a small asymmetry of visuospatial attention that favours the left side of space relative to the right (pseudoneglect). However, it remains unclear whether this leftward bias is maintained, eliminated or shifted rightward in older age. Here we present two meta-analyses that aimed to identify whether adults aged ≥50 years old display a group-level spatial attention bias, as indexed by the line bisection and the landmark tasks. A total of 69 datasets from 65 studies, involving 1654 participants, were analysed. In the meta-analysis of the line bisection task (n = 63), no bias was identified for studies where the mean age was ≥50, but there was a clear leftward bias in a subset where all individual participants were aged ≥50. There was no moderating effect of the participant's age or sex, line length, line position, nor the presence of left or right cues. There was a small publication bias in favour of reporting rightward biases. Of note, biases were slightly more leftward in studies where participants had been recruited as part of a stand-alone older group, compared to studies where participants were recruited as controls for a clinical study. Similarly, no spatial bias was observed in the meta-analysis of the landmark task, although the number of studies included was small (n = 6). Overall, these results indicate that over 50s maintain a group-level leftward bias on the line bisection task, but more studies are needed to determine whether this bias can be modulated by stimulus- or state-dependent factors.

Signatures of functional visuospatial asymmetries in early infancy.

Adults present a large number of asymmetries in visuospatial behavior that are known to be supported by functional brain lateralization. Although there is evidence of lateralization for motor behavior and language processing in infancy, no study has explored visuospatial attention biases in the early stages of development. In this study, we tested for the presence of a leftward visuospatial bias (i.e., pseudoneglect) in 4- and 5-month-old infants using an adapted version of the line bisection task. Infants were trained to identify the center of a horizontal line (Experiment 1) while their eye gazes were monitored using a remote eye-tracking procedure to measure their potential gazing error. Infants exhibited a robust pseudoneglect, gazing leftward with respect to the veridical midpoint of the horizontal line. To investigate whether infants' pseudoneglect generalizes to any given object or is dependent on the horizontal dimension, in Experiment 2 we assessed infants' gaze deployment in vertically oriented lines. No leftward bias was found, suggesting that early visuospatial attention biases in infancy are constrained by the orientation of the visual plane in which the information is organized. The interplay between biological and cultural factors that might contribute to the early establishment of the observed leftward bias in the allocation of visuospatial attention is discussed.

On line bisection: Validity and reliability of online measures of pseudoneglect.

This study assessed pseudoneglect using line bisection and perceptual landmark tasks in two matched online sessions. Line bisection bias was characterized by the traditional measure of Directional Bisection Error (DBE), and by Endpoint Weightings Bias (EWB), derived from an "endpoint weightings" analysis, made possible by the independent manipulation of left and right endpoints. EWB is proposed to index the relative attentional allocation to the two ends of the line. The expected leftward bias (pseudoneglect) was found, with larger effect sizes for EWB (d = -0.34 in both sessions) than for DBE (-0.22 in Session 1 and -0.14 in Session 2). Although EWB was slightly less reliable than DBE, it was more sensitive to pseudoneglect, and the endpoint weightings method has further advantages, including the option of an additional measure of non-lateralized attention. A substantial proportion of participants had difficulty following the instructions for the landmark task, which highlights the need for clear instructions and performance checks for this task. This study shows that line bisection can be used to measure pseudoneglect online, and provides grounds to suggest that the task should routinely include the independent manipulation of left and right endpoints, so that an endpoint weightings analysis can be performed.

Righteous Adam, Sinister Eve.

The symbolism of laterality in images implies that the virtuous figure is represented on the right of the scene whereas the sinful character is depicted on the left. In portraits of male and female characters this has reflected and reinforced stereotypes and inequalities down the ages. Given these premises, we hypothesized that the prototypical representations of Adam and Eve, as a man and a woman conflated with notions of virtue and vice, would show a non-random arrangement. We tested this hypothesis, sampling artistic depictions of the Garden of Eden, from the twelth century to the present day in three separately-collected series of 100, 99, and 142 images respectively. Eve is depicted to Adam's left significantly more often than chance (between 70% and 83%), particularly in pre-1600 artworks. We interpret this asymmetry as reflecting the perceived lesser status of women in relation to men, since the allegorical incipit of humankind. We also provide experimental evidence that this asymmetry, although pervasive, has not been internalized by modern viewers. Cognitive sciences account for this spatial asymmetry in terms of preference for figures placed within the left visual field of the observer, i.e., within the right space of the objective scene.

Spatial asymmetries ("pseudoneglect") in free visual exploration-modulation of age and relationship to line bisection.

When humans visually explore an image, they typically tend to start exploring its left side. This phenomenon, so-called pseudoneglect, is well known, but its time-course has only sparsely been studied. Furthermore, it is unclear whether age influences pseudoneglect, and the relationship between visuo-spatial attentional asymmetries in a free visual exploration task and a classical line bisection task has not been established. To address these questions, 60 healthy participants, aged between 22 and 86, were assessed by means of a free visual exploration task with a series of naturalistic, colour photographs of everyday scenes, while their gaze was recorded by means of a contact-free eye-tracking system. Furthermore, a classical line bisection task was administered, and information concerning handedness and subjective alertness during the experiment was obtained. The results revealed a time-sensitive window during visual exploration, between 260 and 960 ms, in which age was a significant predictor of the leftward bias in gaze position, i.e., of pseudoneglect. Moreover, pseudoneglect as assessed by the line bisection task correlated with the average gaze position throughout a time-window of 300-1490 ms during the visual exploration task. These results suggest that age influences visual exploration and pseudoneglect in a time-sensitive fashion, and that the degree of pseudoneglect in the line bisection task correlates with the average gaze position during visual exploration in a time-sensitive manner.

Disconnected hand avatar can be integrated into the peripersonal space.

Several studies have shown that space immediately surrounding the body, or the peripersonal space is represented differently in the brain from the more distant extra-personal space. Moreover, the boundary of peripersonal space can be extended to space surrounding the tip of a tool held by the hand. However, it is not known if tools need to be connected to the body to modulate the peripersonal space. We used a line bisection task to investigate whether peripersonal space representation surrounds a virtual hand avatar that is disconnected from the body. Healthy participants conducted a line bisection task by responding with either a virtual hand avatar or a laser pointer. The to-be-bisected lines were presented either in peripersonal or extra-personal space. When the lines were placed in extra-personal space, the virtual hand avatar was presented near the line such that the hand avatar was far from participants and disconnected from their bodies. Results indicated a shift in the line bisection bias from the left to the right as the line presentation distance increased when using the laser pointer, whereas no shift in bias was observed when using the virtual hand avatar. This result indicates that objects resembling human hands presented even at a distance and disconnected from the body can be integrated into the peripersonal space, which suggests that peripersonal space representation is more flexible than previously reported.

The effect of cognitive load on horizontal and vertical spatial asymmetries.

Healthy individuals typically show a leftward attentional bias in the allocation of spatial attention along the horizontal plane, a phenomenon known as pseudoneglect, which relies on a right hemispheric dominance for visuospatial processing. Also, healthy individuals tend to overestimate the upper hemispace when orienting attention along the vertical plane, a phenomenon that may depend on asymmetric ventral and dorsal visual streams activation. Previous research has demonstrated that when attentional resources are reduced due to increased cognitive load, pseudoneglect is attenuated (or even reversed), due to decreased right-hemispheric activations. Critically, whether and how the reduction of attentional resources under load modulates vertical spatial asymmetries has not been addressed before. We asked participants to perform a line bisection task both with and without the addition of a concurrent auditory working memory task with lines oriented either horizontally or vertically. Results showed that increasing cognitive load reduced the typical leftward/upward bias with no difference between orientations. Our data suggest that the degree of cognitive load affects spatial attention not only in the horizontal but also in the vertical plane. Lastly, the similar effect of load on horizontal and vertical judgements suggests these biases may be related to only partially independent mechanisms.

Mapping the anatomy of perceptual pseudoneglect. A multivariate approach.

Fundamental to the understanding of the functions of spatial cognition and attention is to clarify the underlying neural mechanisms. It is clear that relatively right-dominant activity in ventral and dorsal parieto-frontal cortex is associated with attentional reorienting, certain forms of mental imagery and spatial working memory for higher loads, while lesions mostly to right ventral areas cause spatial neglect with pathological attentional biases to the right side. In contrast, complementary leftward biases in healthy people, called pseudoneglect, have been associated with varying patterns of cortical activity. Notably, this inconsistency may be explained, at least in part, by the fact that pseudoneglect studies have often employed experimental paradigms that do not control sufficiently for cognitive processes unrelated to pseudoneglect. To address this issue, here we administered a carefully designed continuum of pseudoneglect and control tasks in healthy adults while using functional magnetic resonance imaging (fMRI). Data submitted to partial least square (PLS) imaging analysis yielded a significant latent variable that identified a right-dominant network of brain regions along the intra-occipital and -parietal sulci, frontal eye fields and right ventral cortex in association with perceptual pseudoneglect. Our results shed new light on the interplay of attentional and cognitive systems in pseudoneglect.

The effects of hemispheric dominance, literacy acquisition, and handedness on the development of visuospatial attention: A study in preschoolers and second graders.

A tendency to over-attend the left side of the space (i.e., pseudoneglect) has been repeatedly reported in Western adult populations and is supposed to reflect a right hemisphere dominance in the control of visuospatial attention. This neurobiological hypothesis has been partially challenged by growing evidence showing that pseudoneglect is profoundly triggered by cultural practices such as reading and writing habits. Accordingly, more recent theoretical accounts suggest a strict coupling between nature and nurture dimensions at the origins of such bias. To further explore this possibility, here we first administered a digitized cancellation task to right-handed Western children before and after literacy acquisition. Results showed an incremental leftward shift of attention in the cancellation of the first target and an increasing preference for a left-to-right visual search from preschoolers to second graders. Yet, despite these differences, the overall distribution of visuospatial attention was biased to the left in both groups. To explore the role of handedness in visuospatial asymmetries, we also tested a group of left-handed second graders. Results showed an impact of handedness on visuospatial performance, with an accentuated rightward-oriented visual search for left-handed children, although the overall distribution of attention was again biased to the left hemispace. Taken together, these findings do not provide support to a pure neurobiological view of visuospatial biases. Rather, our study indicates that the control of visuospatial attention is mediated by a dynamic interplay among biological (i.e., right hemisphere dominance), biomechanical (i.e., hand dominance), and cultural (i.e., reading habits) factors.

A leftward bias for the arrangement of consumer items that differ in attractiveness.

People are frequently biased to use left-side information more than right-side information to inform their perceptual judgements. This research examined whether the leftward bias also applied to preferences for the arrangement of everyday consumer items. Pairs of consumer items were created where one item was more attractive than the other item. Using a two-alternative forced choice task, Experiment 1 found a robust preference for arrangements with the more attractive consumer item on the left side rather than the right side of a pair. Experiment 2 reversed the judgement decision, with participants asked to choose the arrangement they least preferred, and a bias for arrangements with the more attractive item on the right side emerged. Experiment 3 failed to find an effect of the "attractive left" preference on participants' purchasing intentions. The preference for attractive left arrangements has implications for the display of consumer products and for the aesthetic arrangement of objects in general. The findings are discussed in relation to hemispheric asymmetries in processing and the role of left to right scanning.

Lateral biases in aesthetic and spatial location judgments: differences between tasks and native reading directions.

People exhibit consistent leftward spatial biases across a variety of tasks. However, individuals with a native reading direction other than left-to-right (LTR) show an attenuation of the leftward bias. The current study used procedurally similar tasks to examine spatial ability and aesthetic preferences in LTR and right-to-left (RTL) groups. In the spatial task participants viewed a centred rectangle partially occluded by an overlapping circle and estimated the centre of the circle with a single mouse click. In the aesthetic task participants used the mouse to control a "virtual flashlight" to light images of abstract paintings in the most aesthetically pleasing way. Contrary to predictions, smaller errors were made for circles on the right and estimations were progressively less accurate as circle size increased in the spatial task. On the aesthetic task, light placements of LTR participants were biased to the left and significantly different from the slightly rightward placements of RTL participants. As predicted, when completing the aesthetics task amounts of time scanning left or right visual space were different between groups. Findings support the theory that directional scanning biases attenuate leftward lateral biases and further, the nature of the visuospatial task may vary the strength of lateral bias.

Improved Accuracy on Lateralized Spatial Judgments in Healthy Aging.

OBJECTIVES: Healthy young adults often demonstrate a leftward spatial bias called "pseudoneglect" which often diminishes with aging. One hypothesis for this phenomenon is an age-related deterioration in right hemisphere functions (right hemi-aging). If true, then a greater rightward bias should be evident on all spatial attention tasks regardless of content. Another hypothesis is a decrease in asymmetrical hemispheric activation with age (HAROLD). If true, older participants may show reduced bias in all spatial tasks, regardless of leftward or rightward biasing of specific spatial content. METHODS: Seventy right-handed healthy participants, 33 younger (21-40) and 37 older (60-78), were asked to bisect solid and character-letter lines as well as to perform left and right trisections of solid lines. RESULTS: Both groups deviated toward the left on solid line bisections and left trisections. Both groups deviated toward the right on right trisections and character line bisections. In all tasks, the older participants were more accurate than the younger participants. CONCLUSIONS: The finding that older participants were more accurate than younger participants across all bisection and trisection conditions suggests a decrease in the asymmetrical hemispheric activation of these specialized networks important in the allocation of contralateral spatial attention or spatial action intention.

The Influence of Focused and Sustained Spatial Attention on the Allocation of Spatial Attention.

OBJECTIVE: The objective of this study was to evaluate the impact of directed and sustained attention on the allocation of visuospatial attention. Healthy people often have left lateral and upward vertical spatial attentional biases. However, it is not known whether there will be an increase in bias toward the attended portion of the stimulus when volitional spatial attention is allocated to a portion of a stimulus, whether there are asymmetrical spatial alterations of these biases, and how sustained attention influences these biases. METHODS: We assessed spatial bias in 36 healthy, right-handed participants using a variant of horizontal and vertical line bisections. Participants were asked to focus on one or the other end of vertical or horizontal lines or entire vertical or horizontal lines, and then to bisect the line either immediately or after a 20 second delay. RESULTS: We found a significant main effect of attentional focus and an interaction between attentional focus and prolonged viewing with delayed bisection. Focusing on a certain portion of the line resulting in a significant deviation toward the attended portion and prolonged viewing of the line prior to bisection significantly enhanced the degree of deviation toward the attended portion. CONCLUSIONS: The enhanced bias with directed and sustained attention may be useful modifications of the line bisection test, particularly in clinical populations. Thus, future studies should determine whether prolonged viewing with delayed bisection and spatially focused attention reveals attentional biases in patients with hemispheric lesions who perform normally on the traditional line bisection test. (JINS, 2019, 25, 65-71).