Biases of spatial attention in vision and audition

Neurologically normal observers misperceive the midpoint of horizontal lines as systematically leftward of veridical center, a phenomenon known as pseudoneglect. Pseudoneglect is attributed to a tonic asymmetry of visuospatial attention favoring left hemispace. Whereas visuospatial attention is biased toward left hemispace, some evidence suggests that audiospatial attention may possess a right hemispatial bias. If spatial attention is supramodal, then the leftward bias observed in visual line bisection should also be expressed in auditory bisection tasks. If spatial attention is modality specific then bisection errors in visual and auditory spatial judgments are potentially dissociable. Subjects performed a bisection task for spatial intervals defined by auditory stimuli, as well as a tachistoscopic visual line bisection task. Subjects showed a significant leftward bias in the visual line bisection task and a significant rightward bias in the auditory interval bisection task. Performance across both tasks was, however, significantly positively correlated. These results imply the existence of both modality specific and supramodal attentional mechanisms where visuospatial attention has a prepotent leftward vector and audiospatial attention has a prepotent rightward vector of attention. In addition, the biases of both visuospatial and audiospatial attention are correlated.     

Concurrent processing of spatial relations and objects in two cerebral hemispheres

This study examined hemispheric asymmetry for concurrent processing of object and spatial information. Participants viewed two successive stimuli, each of which consisted of two digits and two pictures that were randomly located and judged them as identical or different. A sample stimulus was presented in a central visual field, followed by a matching stimulus presented briefly in a left or right visual field. The matching stimuli were different from the sample stimuli with respect to the object (digit or picture) or spatial (locations or distances of items) aspect. No visual field asymmetry was found in the detection of object change. However, a left visual field advantage was found in the detection of spatial change. This result can be explained by the double filtering by frequency theory of Ivry and Robertson, who asserted that the left hemisphere has a bias for processing information contained in relatively high spatial frequencies whereas the right hemisphere has a bias for processing information contained in relatively low spatial frequencies. Based upon this evidence, the importance of interhemispheric integration for visual scene perception is discussed.     

Representational neglect for words as revealed by bisection tasks

In the present study, we showed that a representational disorder for words can dissociate from both representational neglect for objects and neglect dyslexia. This study involved 14 brain-damaged patients with left unilateral spatial neglect and a group of normal subjects. Patients were divided into four groups based on presence of left neglect dyslexia and representational neglect for non-verbal material, as evaluated by the Clock Drawing test. The patients were presented with bisection tasks for words and lines. The word bisection tasks (with words of five and seven letters) comprised the following: (1) representational bisection: the experimenter pronounced a word and then asked the patient to name the letter in the middle position; (2) visual bisection: same as (1) with stimuli presented visually; and (3) motor bisection: the patient was asked to cross out the letter in the middle position. The standard line bisection task was presented using lines of different length. Consistent with the literature, long lines were bisected to the right and short lines, rendered comparable in length to the words of the word bisection test, deviated to the left (crossover effect). Both patients and controls showed the same leftward bias on words in the visual and motor bisection conditions. A significant difference emerged between the groups only in the case of the representational bisection task, whereas the group exhibiting neglect dyslexia associated with representational neglect for objects showed a significant rightward bias, while the other three patient groups and the controls showed a leftward bisection bias. Neither the presence of neglect alone nor the presence of visual neglect dyslexia was sufficient to produce a specific disorder in mental imagery. These results demonstrate a specific representational neglect for words independent of both representational neglect and neglect dyslexia.     

Visuospatial Neglect: The Ultimate Deconstruction?

Unilateral visuospatial neglect is now widely acknowledged to be a highly heterogeneous condition: The overt manifestations of visual neglect can vary as a function of task, spatial domain, and mode of response (at least). Double dissociations (sometimes of the strong form) have already been reported between most of the components of what was originally thought to be a relatively stable construct within the visual modality. Nonetheless, throughout successive fractionations of neglect, reported cases of bidirectional task-specific neglect after unilateral brain damage are rare. We now report two such cases. After right hemisphere stroke, the first patient reliably showed severe left neglect on cancellation but right neglect on line bisection. After left hemisphere stroke, the second case showed right neglect on cancellation but left neglect on line bisection. Extensive investigation of case 1 confirmed our previous conjecture that the crucial distinction between these tasks lies in the presence or the absence of an overt target. In contrast to cancellation, line bisection demands the internal computation of the location of the “target” (the midpoint), followed by executing a motor response toward the precise location of that “imaginary” midpoint. The relative attentional and premotor contributions of the intact and damaged hemispheres to these forms of bidirectional neglect are also assessed.     

The relationship between vertical stimulation and horizontal attentional asymmetries

The original aim was to examine the effect of perceived distance, induced by the Ponzo illusion, on left/right asymmetries for line bisection. In Experiment 1, university students (n?=?29) made left/right bisection judgements for lines presented in the lower or upper half of the screen against backgrounds of the Ponzo stimuli, or a baseline. While the Ponzo illusion had relatively little effect on line bisection, elevation in the baseline condition had a strong effect, whereby the leftward bias was increased for upper lines. Experiment 2 (n?=?17) eliminated the effect of elevation by presenting the line in the middle and moving the Ponzo stimuli relative to the line. Despite this change, the leftward bias was still stronger in the upper condition in the baseline condition. The final experiment (n?=?17) investigated whether upper/lower visual stimulation, which was irrelevant to the task, affected asymmetries for line bisection. The results revealed that a rectangle presented in the upper half of the screen increased the leftward line bisection bias relative to a baseline and lower stimulation condition. These results corroborate neuroimaging research, showing increased right parietal activation associated with shifts of attention into the upper hemispace. This increased right parietal activation may increase the leftward attentional bias—resulting in a stronger leftward bias for line bisection.     

Understanding the effects of moving visual stimuli on unilateral neglect following stroke

Moving visual stimuli have been shown to reduce unilateral neglect (ULN), however, the mechanisms underlying these effects remain poorly understood. This study compared lateralised and non-lateralised moving visual stimuli to investigate whether the spatial characteristics or general alerting properties of moving visual stimuli are responsible for reducing neglect. Post-stroke left neglect patients as well as healthy and patient control subjects were tested on a computerised line bisection task under six visual stimulus conditions. The key finding was that, relative to the no stimulus condition, leftward moving and left-sided moving visual stimuli shifted neglect patients' bisection errors leftward while the non-lateralised random moving visual stimuli did not reduce neglect patients' rightward bisection errors. The results provide evidence that spatial characteristics rather than general alerting properties of moving visual stimuli reduce rightward bisection errors in ULN. Moreover, the pattern of findings strongly supports the notion that moving visual stimuli reduce neglect by capturing attention and drawing it to a spatial location rather than by activating the attentional system via superior collicular neurons.     

Radial bisection of words and lines in right‐brain‐damaged patients with spatial neglect

The bisection of lines positioned radially (with the two ends of the line close and far, with respect to the participant's body) has been less investigated than that of lines placed horizontally (with their two ends left and right, with respect to the body's midsagittal plane). In horizontal bisection, patients with left neglect typically show a rightward bias for both lines and words, greater with longer stimuli. As for radial bisection, available data indicate that neurologically unimpaired participants make a distal error, while results from right‐brain‐damaged patients with left spatial neglect are contradictory. We investigated the bisection of radially oriented words, with the prediction that, during bisection, linguistic material would be recoded to its canonical left‐to‐right format in reading, with the performance of neglect patients being similar to that for horizontal words. Thirteen right‐brain‐damaged patients (seven with left spatial neglect) and fourteen healthy controls were asked to manually bisect 40 radial and 40 horizontal words (5–10 letters), and 80 lines, 40 radial and 40 horizontal, of comparable length. Right‐brain‐damaged patients with spatial neglect exhibited a proximal bias in the bisection of short radial words, with the proximal part corresponding to the final right part of horizontally oriented words. This proximal error was not found in patients without neglect and healthy controls. For bisection, short radial words may be recoded to the canonical orthographic horizontal format, unveiling the impact of left neglect on radially oriented stimuli.     

Do working memory-driven attention shifts speed up visual awareness?

Previous research has shown that content representations in working memory (WM) can bias attention in favor of matching stimuli in the scene. Using a visual prior-entry procedure, we here investigate whether such WM-driven attention shifts can speed up the conscious awareness of memory-matching relative to memory-mismatching stimuli. Participants were asked to hold a color cue in WM and to subsequently perform a temporal order judgment (TOJ) task by reporting either of two different-colored circles (presented to the left and right of fixation with a variable temporal interval) as having the first onset. One of the two TOJ circles could match the memory cue in color. We found that awareness of the temporal order of the circle onsets was not affected by the contents of WM, even when participants were explicitly informed that one of the TOJ circles would always match the WM contents. The null effect of WM on TOJs was not due to an inability of the memory-matching item to capture attention, since response times to the target in a follow-up experiment were improved when it appeared at the location of the memory-matching item. The present findings suggest that WM-driven attention shifts cannot accelerate phenomenal awareness of matching stimuli in the visual field.     

The visual basis of category effects in object identification: evidence from the visual hemifield paradigm

The basis for the category specific living things advantage in object recognition (i.e., faster and more accurate identification of living compared to nonliving things) was investigated in two experiments. It was hypothesised that the global shape of living things on average provides more information about their basic level identity than the global shape of nonliving things. In two experiments subjects performed name-picture or picture-name verification tasks, in which blurred or clear images of living and nonliving things were presented in either the right or the left visual hemifield. With blurred images, recognition performance was worst for nonliving things presented to the right visual field/left hemisphere, indicating that the lack of visual detail in the stimulus combined with a left hemisphere bias toward processing high frequency visual elements proved detrimental for processing nonliving stimuli in this condition. In addition, an overall living things advantage was observed in both experiments. This advantage was considerably larger with blurred images than with clear. These results are compatible with the global shape hypothesis and converge with evidence using other paradigms.     

Virtual reality pencil and paper tests for neglect: a protocol.

Hemispatial neglect is a common disorder that can occur after a lesion in the right hemisphere. Its main characterization is the difficulty in processing visual stimuli emanating from the space contralateral to the lesion. Pencil and paper tests (such as target cancellation, line bisection, or drawing copy) are used to diagnose neglect. We propose using virtual reality technology and haptic force feedback to enhance pencil and paper tests. Our system can track the patients' eye-gaze and their hand movements. Also, the efficiency of several techniques used to decrease the neglect in different sensory spaces are investigated.     

A Search for the Optimal Stimulus

How do stimulus size and item number relate to the magnitude and direction of error on center estimation and line cancellation tests? How might this relationship inform theories concerning spatial neglect? These questions were addressed by testing twenty patients with right hemisphere lesions, eleven with left hemisphere lesions and eleven normal control subjects on multiple versions of center estimation and line cancellation tests. Patients who made large errors on these tests also demonstrated an optimal or pivotal stimulus value, i.e., a particular size center estimation test or number of lines on cancellation that either minimized error magnitude relative to other size stimuli (optimal) or marked the boundary between normal and abnormal performance (pivotal). Patients with right hemisphere lesions made increasingly greater errors on the center estimation test as stimuli were both larger and smaller than the optimal value, whereas those with left hemisphere lesions made greater errors as stimuli were smaller than a pivotal value. In normal subjects, the direction of errors on center estimation stimuli shifted from the right of true center to the left as stimuli decreased in size (i.e., the crossover effect). Right hemisphere lesions exaggerated this effect, whereas left hemisphere lesions diminished and possibly reversed the direction of crossover. Error direction did not change as a function of stimulus value on cancellation tests. The demonstration of optimal and pivotal stimulus values indicates that performances on center estimation and cancellation tests in neglect are only relative to the stimuli used. In light of other studies, our findings indicate that patients with spatial neglect grossly overestimate the size of small stimuli and underestimate the size of large stimuli, that crossover represents an “apparent” shift in error direction that actually results from normally occurring errors in size perception, and that the left hemisphere is specialized for one aspect of size estimation, whereas the right performs dual roles.     

Perceptual asymmetries in normal children and children with attention deficit/hyperactivity disorder

Perceptual asymmetries in normal right-handed children (7-12 years of age) and children with attention deficit/hyperactivity disorder (ADHD), combined type, were investigated using various chimeric stimuli in free-viewing conditions. In the face-matching task, participants indicated which of two symmetrical face chimeras more closely resembled the original; in the chimeric faces task, participants indicated which of a pair of faces appeared happier; and in the grey scales task participants indicated which of two shaded rectangles appeared to be darker overall. Leftward biases were found for normal children with no effects of age. Contrary to expectations, normal leftward biases were also found for ADHD children in the face-matching and the chimeric faces tasks; however, a significant leftward bias was not observed in the grey scales task. The absence of anomalous perceptual bias in ADHD children on these purely perceptual tasks, suggests that anomalous perceptual asymmetries observed in other tasks (line bisection and cancellation tasks) may have been confounded by the motor response, and/or the explicit spatial components of those tasks.     

Auditory space perception in left- and right-handers

Several studies have shown that handedness has an impact on visual spatial abilities. Here we investigated the effect of laterality on auditory space perception. Participants (33 right-handers, 20 left-handers) completed two tasks of sound localization. In a dark, anechoic, and sound-proof room, sound stimuli (broadband noise) were presented via 21 loudspeakers mounted horizontally (from 80° on the left to 80° on the right). Participants had to localize the target either by using a swivel hand-pointer or by head-pointing. Individual lateral preferences of eye, ear, hand, and foot were obtained using a questionnaire. With both pointing methods, participants showed a bias in sound localization that was to the side contralateral to the preferred hand, an effect that was unrelated to their overall precision. This partially parallels findings in the visual modality as left-handers typically have a more rightward bias in visual line bisection compared with right-handers. Despite the differences in neural processing of auditory and visual spatial information these findings show similar effects of lateral preference on auditory and visual spatial perception. This suggests that supramodal neural processes are involved in the mechanisms generating laterality in space perception.     

Blind individuals show pseudoneglect in bisecting numerical intervals

Neurologically normal individuals typically show a leftward bias--known as pseudoneglect--in bisecting physical lines as well as numerical intervals. The latter bias may reflect the spatial nature in which numbers are represented (i.e., the mental number line). In previous studies, we found that congenitally blind individuals show such leftward bias in haptic bisection. Here, we demonstrate that blind individuals also show a consistent leftward bias in numerical bisection. The leftward bias was greater when numbers were presented in descending rather than ascending order, and the magnitude of the leftward bias was comparable to that shown by a control group of blindfolded sighted participants. Our findings thus support the view that pseudoneglect operates at a mental representational level rather than being perceptually based. Moreover, the consistent leftward bias shown by blind individuals in both line and numerical bisection suggests that the right hemisphere dominance in spatial processing, resulting in an overestimation of the left side of space, develops even in the absence of any visual input.     

Number line estimation and complex mental calculation: Is there a shared cognitive process driving the two tasks?

It is widely accepted that different number-related tasks, including solving simple addition and subtraction, may induce attentional shifts on the so-called mental number line, which represents larger numbers on the right and smaller numbers on the left. Recently, it has been shown that different number-related tasks also employ spatial attention shifts along with general cognitive processes. Here we investigated for the first time whether number line estimation and complex mental arithmetic recruit a common mechanism in healthy adults. Participants’ performance in two-digit mental additions and subtractions using visual stimuli was compared with their performance in a mental bisection task using auditory numerical intervals. Results showed significant correlations between participants’ performance in number line bisection and that in two-digit mental arithmetic operations, especially in additions, providing a first proof of a shared cognitive mechanism (or multiple shared cognitive mechanisms) between auditory number bisection and complex mental calculation.     

The modulation of haptic line bisection by a visual illusion and optokinetic stimulation

Research has shown that a variety of different sensory manipulations, including visual illusions, transcutaneous nerve stimulation, vestibular caloric stimulation, optokinetic stimulation, and prism adaptation, can all influence people's performance on spatial tasks such as line bisection. It has been suggested that these manipulations may act upon the 'higher-order' levels of representation used to code spatial information. We investigated whether we could influence haptic line bisection in normal participants crossmodally by varying the visual background that participants viewed. In experiment 1, participants haptically bisected wooden rods while looking at a variant of the Oppel - Kundt visual illusion. Haptic-bisection judgments were influenced by the orientation of the visual illusion (in line with previous unimodal visual findings). In experiment 2, haptic-bisection judgments were also influenced by the presence of a leftward or rightward moving visual background. In experiments 3 and 4, the position of the to-be-bisected stimuli was varied with respect to the participant's body midline. The results confirmed an effect of optokinetic stimulation, but not of the Oppel -Kundt illusion, on participants' tactile-bisection errors, suggesting that the two manipulations might differentially affect haptic processing. Taken together, these results suggest that the 'higher-order' levels of spatial representation upon which perceptual judgments and/or motor responses are made may have multisensory or amodal characteristics.     

Visual-perceptual abilities in healthy controls, depressed patients, and schizophrenia patients

Previous studies have suggested a right hemineglect in schizophrenia, however few assessed possible visual-perceptual implication in this lateralized anomaly. A manual line bisection without (i.e., lines presented on their own) or with a local cueing paradigm (i.e., a number placed at one or both ends of the line) and the Motor-free Visual Perceptual Test-Vertical format (MVPT-V) were used to assess the visual-perceptual abilities of healthy controls, schizophrenia and depressed patients. Whereas healthy controls and depressed patients showed a non-significant leftward bias in manual line bisection, schizophrenia patients bisected significantly to the left of the true centre of the line. Interestingly, the pattern of performances in response to the local cueing paradigm was similar in depressed and schizophrenia patients such that both groups demonstrated a significant change in their bisection performance only in response to a cue placed at the right extremity of the line (control performance was modified by cues at either end of the line). Finally, in the MVPT-V, schizophrenia patients were impaired relative to the other two groups, especially in the spatial working memory and visual closure categories. These results suggest that: 1/a deficit towards the right hemifield, consistent with a mild form of right hemineglect, can be observed in schizophrenia; 2/lateralized anomalies could also be observed in depression using an appropriate tool such as manual line bisection; 3/performances in the MVPT-V suggested that a simple visual-perceptual deficit could not explain the lateralized anomaly observed in the manual line bisection, as it is the case in the hemineglect syndrome.     

Attentional bias toward low-intensity stimuli: an explanation for the intensity dissociation between reaction time and temporal order judgment?

If two stimuli need different times to be processed, this difference should in principle be reflected both by response times (RT) and by judgments of their temporal order (TOJ). However, several dissociations have been reported between RT and TOJ, e.g., RT is more affected than TOJ when stimulus intensity decreases. One account for these dissociations is to assume differences in the allocation of attention induced by the two tasks. To test this hypothesis, different distributions of attention were induced in the present study between two stimulus positions (above and below fixation). Only bright stimuli appeared in one position and either bright or dim stimuli in the other. In the two RT experiments, participants had to respond to every stimulus appearing in one of the two positions. Reaction times to bright stimuli were faster when they appeared in the position where dim stimuli were likely to occur. This finding suggests that the allocation of attention was adapted to the asymmetrical arrangement of stimuli, not suggested by explicit instruction. In the two TOJ experiments, the temporal order of stimuli appearing in the two positions had to be judged. Although bright stimuli appearing at the bright-and-dim location were judged to be earlier, this effect was small and insignificant. Further, the intensity dissociation between RT and TOJ was insensitive to random vs blockwise presentations of intensities, therefore was not modified by attentional preferences. Thus, asymmetrical arrangement of stimuli has an impact on the allocation of attention, but only in the RT task. Therefore dissociations between TOJ and response times cannot be accounted for by an attentional bias in the TOJ task but probably by different use of temporal information in the two tasks.     

Using scissors to bisect a line: A perception–action dissociation in complex tool use

Previous research has shown that unilateral visual neglect is improved when patients are required to actively grasp an object at its center, rather than only pointing to its center. A similar dissociation between pointing and grasping responses has been reported for pseudoneglect, a spatial bias toward the left side of space that is normally exhibited by healthy participants. Among other theories, the two-visual-streams hypothesis has been offered as an explanation for these dissociations. According to this hypothesis, highly practiced actions that are performed under easily available visual input (e.g., grasping an object) are controlled by the dorsal stream and can resist perceptual judgment biases. We investigated whether, in pseudoneglect, dissociations between perceptual judgments and motor–manipulative responses occur when participants have to respond with complex tools that are highly practiced but that have been shown to rely on both ventral- and dorsal-stream processing. In a standard line bisection task, participants had to either mark the center of a line with a pencil or cut the line in two halves using a pair of scissors. The results indicated the typical leftward bias (pseudoneglect) in the pencil task, whereas performance was much more accurate in the scissors task. These results show that actions featuring complex tool use can resist perceptual judgment biases, and the findings are discussed with reference to existing accounts of perception–action dissociations.