The eyes have it: Oculomotor exploration and line bisection in neglect

In this issue of Cortex, Ishiai et al. (2006) report the eye movements of patients with left neglect during the bisection of lines of different lengths. This is the latest in a series of papers from Ishiai's group, which form an important corpus of data on the oculomotor behaviour of neglect patients during line bisection and related tasks. In this article, I argue that these data should critically constrain theoretical models of bisection errors in neglect, but that these constraints have been applied rarely in practice. First, I briefly introduce bisection behaviour in neglect and describe some of the models proposed to account for its character. I then consider the implications for these models of Ishiai and colleagues' observations. Finally, I outline a novel view of the bisection task that is more compatible with their observations.     

Orientation bias in unilateral neglect: representational contributions

Left-neglect patients bisect horizontal lines to the right of true center. Longer lines are bisected further to the right than shorter lines. This line-length effect might be explained by an increase in the rightward bias of attention because longer lines extend further ipsilesionally. Alternatively, neglect patients might be limited in their abilities to internally represent horizontal magnitudes. Patients might orient further rightward with longer lines because these lines have longer representations. If the line-length effect occurs on lines of identical objective length but they are represented differently, then central mechanisms must contribute to the orientation bias. We constructed two types of lines that were perceived by normal subjects as having different lengths, but were of identical extents. Three neglect patients bisected lines perceived as longer, further to the right than lines perceived as shorter. These results demonstrate that relative magnitudes of internal representations contribute to the degree of bias in neglect patients.     

Effects of cueing on visuospatial processing in unilateral spatial neglect

Patients with typical left unilateral spatial neglect bisected lines after cueing to the left end-point, the fixation point being monitored with an eye camera. They persisted with the point of initial fixation made after cueing and placed the mark there without searching leftwards again. The rightward shift of fixation to the initial point of fixation thus determined the location of the subjective midpoint. We consider that rightward attentional bias increased the amplitude of this shift that was planned on the basis of the perception of the whole line while cueing. This hypothesis may explain smaller but obvious rightward bisection errors found in the cueing condition.     

Acceleration perception and spatial distortion in a left unilateral neglect patient

To explain relative leftward overextension in a line extension task by left unilateral neglect subjects, Bisiach et al. (1998) suggested that the representation of space is distorted--i.e., dilated towards the left side. If perception of the velocity of a moving stimulus is due to a calculation of the distance covered per unit time in representational space, then a stimulus with uniform linear motion should be perceived as decelerating when moving leftwards in the visual field of a subject with left unilateral neglect. We investigated the perception of acceleration in a patient with left unilateral neglect and spatial distortion (revealed as relative left overextension in a line extension task) using a task in which the stimuli were right and left moving targets with variable acceleration. The patient's ability to perceive acceleration was much lower (higher acceleration threshold) for leftward movements than rightward movements. Fourteen months later unilateral neglect had improved, and the relative left overextension and decreasing acceleration threshold for leftward movements were reduced. By contrast, alterations in the perception of acceleration for leftward movements were not found in a patient with left unilateral neglect and left underextension and in a patient with right brain damage and left hemianopia. These findings in one patient with left spatial unilateral neglect and a relative left overextension in a line extension task are consistent with the hypothesis that representational space is distorted, with a disproportionate leftward expansion, that affects perception of movement.     

Attention biases the perceived midpoint of horizontal lines

In patients with right brain damage and left visual neglect, attention tends to be captured by right-sided objects and cannot easily disengage from them. While these phenomena can account for several clinical and experimental patterns of performance such as biased visual search, its role is more controversial for other neglect-related signs, such as the typical rightward shifts in horizontal line bisection. It is thus important to see whether and how attentional orienting can bias line bisection in normal participants using standard clinical bisection stimuli. In 3 experiments, we explored the Attentional Repulsion Effect (ARE, Suzuki & Cavanagh, 1997) on pre-bisected lines. Normal observers saw horizontal lines with a vertical bisection mark near the center, preceded by a cue to the left or right of the line, or by no cue. On each trial, observers indicated whether they saw the bisection mark to the left or at the right of the midpoint. We plotted the proportion of ‘seen-at-right’ responses as a function of the mark's actual position. For uncued lines, the point of subjective equality was slightly at the left of the true center, consistent with the pseudoneglect phenomenon. Right-sided cues shifted the apparent bisection point to the left (and vice versa), as predicted by the ARE. Similar results occurred with different task instructions (compare the length of the left-sided line segment to the right-sided segment) and in the presence or absence of central fixation marks. These results obtained in normal participants support attentional accounts of biased line bisection in neglect patients.     

Unilateral spatial neglect due to hemorrhage in the thalamic region

In a series of 33 cases of thalamic hemorrhage, unilateral spatial neglect (USN) was found in 11 patients all of whom had right-sided lesions. An analysis of results suggests that the posteromedial portion of the thalamus plays an important role in the pathogenesis of USN and that extensive lesions beyond the thalamus involving nearby structures may be necessary to cause persistent USN.     

Ineffective leftward search in line bisection and mechanisms of left unilateral spatial neglect

We examined the eye-fixation pattern of a patient with severe left unilateral spatial neglect who showed leftward searches of various extent in more than half of line bisection trials. Because of complete left homonymous hemianopia, he perceived only the segment of the line between its right endpoint and the point of the leftmost fixation. In the trials with leftward searches, he frequently placed the subjective midpoint on the right part of the perceived segment. In the trials without leftward searches, he placed it near the left extreme point of the perceived segment. For all these bisections, the subjective midpoint was constantly placed far to the right of the true midpoint of the line irrespective of the length perceived. We consider that in severe left unilateral spatial neglect, rightward attentional bias is the predominant factor that determines where to place the subjective midpoint. Transient attentional shift to the left may produce leftward searches, but it does not induce effective processing of line bisection.     

Unilateral neglect is not unilateral: evidence for additional neglect of extreme right space

Six patients with visuospatial neglect following right hemisphere lesions were given three tasks that assessed performance in areas of space ranging from extreme left to extreme right. A line bisection task required the patients to detect and bisect lines of four different lengths at seven left-right spatial locations, a number report task required the patients to name 11 two-digit numbers in a left-right array, and a tiling task required patients to place small black tiles over the black squares of a grid that stretched from 65 degrees left to 65 degrees right. Performance was compared with that of 20 age-matched controls. The patients showed the characteristic signs of left-side neglect in left space, extending to the central midline. Performance was relatively normal in centre-right space but all 6 patients showed signs of neglect of extreme right space (60 degrees to the right of the midline and beyond). We propose that neglect is best characterised as a bilateral, asymmetrical compression of experienced space in which the constriction extends further from the left than from the right but nevertheless affects both sides of space.     

Unilateral spatial neglect in Alzheimer's disease A line bisection study

Neuropsychological studies on Alzheimer's disease (AD) have rarely mentioned about unilateral spatial neglect in spite of widespread use of visuospatial tasks. We reported a 62-year-old woman with probable AD who showed moderate dementia with left unilateral spatial neglect and relatively preserved language function. An extensive line bisection study with either hand confirmed her having left unilateral spatial neglect. Single photon emission computed tomography revealed relative hypoperfusion in the right temporal and parietal regions. AD patients with disproportionate right hemisphere dysfunction may exhibit left unilateral spatial neglect if tested adequately in the stage of mild to moderate dementia. We consider that application of the line bisection test to AD patients contributes to estimation of their right hemisphere function.     

Hypermnesia in unilateral neglect

Thirteen right brain-damaged patients who were found to neglect pictures presented on the left of a display were presented the same stimuli, intermixed with foils, in a yes-no recognition test. Fifty per cent of patients claimed to have already seen one or more of the previously neglected pictures. This demonstrates that visual information that fails to access consciousness in neglect patients does retain the ability to surface as explicit memory at a later stage.     

Reading direction and spatial neglect

Many American and European investigators have reported that hemispatial neglect is more frequent and more severe after right than left hemisphere lesions. This hemispheric asymmetry may be due to biological asymmetries, learned behavior, or both. Readers of European languages, unlike readers of Semitic languages, scan from left to right. Learned rightward scanning may increase the unilateral neglect associated with right hemisphere lesions and reduce the severity of neglect associated with left hemisphere lesions. To learn if hemispheric asymmetries of neglect are influenced by learned scanning behavior, we used line bisection and cancellation tasks to study patients with unilateral stroke who read only a Semitic or European language before the age of fifteen. We found that independent of reading direction, unilateral neglect was more commonly associated with right than left hemisphere lesions. After right hemisphere damage right to left readers bisected lines closer to center than left to right readers, but on the cancellation test readers of European languages did not perform differently than readers of Semitic languages. These findings suggest that whereas learned scan direction may influence the severity of neglect when measured by line bisection, these learned directional scans cannot fully account for the observed hemisphere asymmetries of neglect. They also suggest that the line bisection test is more influenced by the direction of scanning than is the cancellation test.     

Dissociated neglect for objective and subjective sizes

We examined the effect of line length and viewing distance on the line bisection performance in near space in five patients with left unilateral spatial neglect following right parietal lesions. A line bisection task by fixation was devised to avoid the influence of manual responses. The rightward deviation measured in visual angle increased linearly as a function of the visual angle of lines 150 mm or more long. This linearity, however, did not hold for lines of 100 mm or less. The deviation measured in length was nearly constant for each of these short lengths, even when the visual angle was varied at different viewing distances. The patients therefore discriminated the objective lengths of the short lines. For small objects, neglect patients may distribute attention mainly on the coordinates scaled for objective size. Received: 16 May 1996 Received in revised form: 1 August 1997 Accepted: 12 August 1997     

Contribution of the superior colliculi to post-stroke unilateral spatial neglect and recovery

Unilateral spatial neglect (USN) is a highly prevalent and disabling consequence of stroke that often responds poorly to existing interventions. Its underlying neural mechanisms are still unclear. Recent work suggests that post-stroke USN may be partly related to a disruption of top-down and bottom-up control of visual attention mediated in part through the midbrain superior colliculi (SC). With mounting evidence from animal and human research, our objectives were: (1) to synthesize the literature implicating the SC in USN; (2) to review the rationale and potential for using eye patching and prism adaptation as USN treatment approaches that recruit SC involvement; and (3) to provide recommendations for research on the potential of therapeutic interventions that involve and/or target the retino-collicular pathway. Given the paucity of human studies, the contribution of the SC in USN, while plausible, remains to be confirmed. Further exploration of the mechanisms involved and their impact on USN in human subjects will help develop theoretically based intervention strategies tailored to USN type.     

A study of right unilateral spatial neglect in left hemispheric lesions: the difference between right-handed and non-right-handed post-stroke patients

We report 20 cases of right unilateral spatial neglect caused by lesions in the left cerebral hemisphere. Differences in neuropsychological symptoms and lesions sites are discussed in connection with handedness. Of the right-handed patients, 6 had severe aphasia, 4 had Gerstmann's syndrome, and 1 had pure agraphia, but unilateral spatial neglect in these cases disappeared after a number of months. Six of the non-right-handed patients had moderate-to-severe aphasia, while the other 3 cases had no aphasia at all. Eight of the 9 cases in this group continued to have right unilateral spatial neglect for more than 6 months. Lesion site as determined by CT differed as to hemisphere, but all fell into the common area previously mentioned in connection with such disorders: i.e., the temporal, parietal and occipital lobes.     

Diverse patterns of performance in copying and severity of unilateral spatial neglect

We analysed the performances of 69 patients with left unilateral spatial neglect in copying a multi-object figure, as well as their performance of the line cancellation and line bisection tests. Three patterns of copying were mainly observed. The first pattern was almost satisfactory performance in copying the central figure with occasional omission of the elements located more laterally to the left. The second pattern was omission of the left half of the whole figure, which was the pattern expected of patients with typical left unilateral spatial neglect. The third pattern was left unilateral spatial neglect when copying the central figure, with favourable copying of left lateral objects; this may be regarded as object-centred neglect. Performance of the line cancellation test, which requires exploratory ability, was significantly better for the patients with the third pattern than for those with the second pattern. A significant difference was also observed between the two groups of patients in performance of the line bisection test, in which exploratory ability plays a more minor role. No significant difference was found in performance of the line cancellation and line bisection tests between the patients with the third and first patterns, in spite of remarkable differences in their copying performances. No distinctive characteristics were found for any group of patients as regards lesion, age, duration after onset, education, and WAJS scores. Therefore we concluded that in most of our patients with neglect, these diverse patterns of copying deficits resulted from different levels of neglect severity but not from different types of unilateral spatial neglect.     

Is unilateral spatial neglect a single phenomenon?

The aim of this study is to report the preliminary findings of a traditional battery of tests and our original battery capable of assessing the presence of components and extent of lesions in patients with unilateral spatial neglect. Thirty patients who had unilateral spatial neglect with a stroke in the right hemisphere were assessed for unilateral spatial neglect on exploratory-motor (E-M) tasks, visual-counting (V-C) tasks, and traditional neglect batteries at least 4 weeks after the onset. Other neuropsychological tests and computed tomography were also performed to investigate the relationship with neglect. A factor analysis showed that our tasks loaded significantly on three factors. E-M neglect was found in 16 patients, and V-C neglect in 22 patients with unilateral spatial neglect. There were high correlations between E-M neglect and motor paralysis, word fluency, backward digit span and motor impersistence. There were high correlations between V-C neglect and visual-field defect, line bisection, line cancellation and figure copying. Lesions in the frontal lobe, caudate, insula, and anterior portion of the paraventricular white matter were commonly associated with E-M neglect. Lesions in the occipital lobe were also associated with V-C neglect. We suggest that unilateral neglect is not a single phenomenon, but rather involves several different components. We propose that E-M and V-C tasks are useful methods for evaluating the extent of lesions in patients with unilateral spatial neglect. Received: 18 November 1996 Received in revised form: 14 March 1997 Accepted: 6 April 1997     

From unilateral neglect to the brain basis of consciousness

According to the dominant focus model, the differential activation of the thalamocortical network is a sufficient condition for the diversity of states of consciousness to occur, and coherent patterns of interactive forebrain activation peaks are sufficient to generate specific conscious figures against ground. These minimal assumptions are consonant with the known properties of unilateral neglect. It becomes unnecessary to suppose that consciousness is an independent entity, occupies a dedicated location in the brain, is contingent on a minimal level of neural complexity, plays a causal role in enabling perception or action, or evolved because it plays any such role.     

Lateral and radial neglect as a function of spatial position: A case study

We report a case of severe visuo-spatial neglect after multiple right-hemisphere infarcts. Experiment 1 demonstrated that the magnitude of transection displacement for the bisection of horizontal lines could be modulated by the lateral position of those lines within peripersonal space. As found in previous studies, neglect was attenuated when the true centre of the lines was to the right of the patient's midsagittal plane. In Experiment 2, radial lines were presented in left, middle, and right space. All bisections were placed too far from the body, but the magnitude of this effect increased significantly from right to left space. In Experiment 3, horizontal lines were presented at three distances within peripersonal space; there was a small but reliable reduction of bisection error for the most distally-placed stimuli. These results suggest that unilateral right hemisphere damage can distort the representation of space (and action therein) more extensively than is implied by the term left neglect.     

"Mind the gap": the size-distance dissociation in visual neglect is a cueing effect

There is a growing body of evidence that the processes mediating the allocation of spatial attention within objects may be separable from those governing attentional distribution between objects. In the neglect literature, a related proposal has been made regarding the perception of (within-object) sizes and (between-object) distances. This proposal follows observations that, in size-matching and bisection tasks, neglect is more strongly expressed when patients are required to attend to the sizes of discrete objects than to the (unfilled) distances between objects. These findings are consistent with a partial dissociation between size and distance processing, but a simpler alternative must also be considered. Whilst a neglect patient may fail to explore the full extent of a solid stimulus, the estimation of an unfilled distance requires that both endpoints be inspected before the task can be attempted at all. The attentional cueing implicit in distance estimation tasks might thus account for their superior performance by neglect patients. We report two bisection studies that address this issue. The first confirmed, amongst patients with left visual neglect, a reliable reduction of rightward error for unfilled "gap" stimuli as compared with solid lines. The second study assessed the cause of this reduction, deconfounding the effects of stimulus type (lines vs. gaps) and attentional cueing, by applying an explicit cueing manipulation to line and gap bisection tasks. Under these matched cueing conditions, all patients performed similarly on line and gap bisection tasks, suggesting that the reduction of neglect typically observed for gap stimuli may be attributable entirely to cueing effects. We found no evidence that a spatial extent, once fully attended, is judged any differently according to whether it is filled or unfilled.     

Straight-ahead pointing correlates with long-line bisection in neglect patients

The spatial bias of neglect patients has been assumed to result from the alteration of a body-centred representation which underlies actions within personal and extra-personal spaces. This deficit seems to correspond to the ipsilesional deviation of the subjective straight-ahead (SSA), often described in neglect patients. However, several recent studies have failed to show significant correlations between the SSA deviation and performance in standard visuo-spatial tests of neglect. Since methodological choices could explain these negative results, the present study re-investigated the question. SSA, assessed by manual pointing, was anchored by instructions to a precise median body part. A body-centred line bisection task was also carried out, during the same session and in similar experimental conditions. Sixteen right brain-damaged patients with (n=8) or without (n=8) neglect, and 8 control subjects free of any neurological problem were included. Taken as a group, the neglect group stood out from the others in both tasks. Moreover, in neglect patients only, SSA position significantly correlated with bisection error for long lines, even if dissociations between SSA shift and bisection error were found in two cases. Results are discussed with regard to the nature of the tasks proposed in the different studies and to methodological aspects that could at least partly explain the aforementioned failures.