Bilateral parietal lesions disrupt the beneficial effects of prism adaptation: evidence from a patient with optic ataxia

Prism adaptation (PA) alleviates some neglect symptoms, however, the mechanisms underlying these effects are unclear. One brain area that may be important in generating these beneficial effects is the superior parietal lobe (SPL), a region not typically damaged in neglect, and known to be important for attention, visuomotor control, and eye movements. We examined the effects of rightward PA on covert attention in CF, a patient with bilateral SPL lesions, compared to a group of controls (N = 26) who underwent sham adaptation. In contrast to previous work in neglect, there was no reduction in CF’s leftward disengage deficit, or rightward attentional bias following PA. These results suggest that the SPL plays an important role in generating the beneficial after-effects of prisms on attention. Both Laure Pisella and James Danckert have contributed equally to this work.     

The disengage deficit in hemispatial neglect is restricted to between-object shifts and is abolished by prism adaptation

We sought to determine the effects of prism adaptation on peripherally cued visual attention shifting in patients with spatial neglect, using a task devised by Egly et al. (J Exp Psychol Gen 123:161–177, 1994) based on the classic Posner paradigm. This task allowed a comparison of “within-object” versus “between-object” attention shifts. A display was presented containing two parallel outline rectangles, and subjects were asked to make rapid responses to a target, which would appear at one end of one of the rectangles. The target location was pre-cued with 75% validity: on invalid trials attention was directed either to the other end of the same rectangle, or to the other rectangle. Healthy subjects and right-hemisphere patients without neglect showed a left-right symmetrical pattern, with a larger validity effect when required to shift attention between rectangles, thus indicating a greater difficulty of attention-shifting between than within the respective shapes. The neglect patients showed the typical leftward “disengage deficit” previously observed in neglect, but only for attention shifts between objects, indicating that the effect is object-based rather than purely spatial. A comparison of vertical and horizontal shift costs showed that this attention-shifting deficit for left-hemifield target stimuli was directional rather than hemifield-based: it was absent for vertical shifts of attention within the left hemifield. Finally, we found that prism adaptation abolished the disengage deficit. We found no effects of prism adaptation in the control subjects. We argue that prism adaptation has a powerful effect on one of the fundamental manifestations of the neglect syndrome. Special issue in honour of Professor G. Berlucchi.     

Left visual neglect: is the disengage deficit space- or object-based?

Attention can be directed to spatial locations or to objects in space. Patients with left unilateral spatial neglect are slow to respond to a left-sided target when it is preceded by a right-sided “invalid” cue, particularly at short cue-target intervals, suggesting an impairment in disengaging attention from the right side in order to orient it leftward. We wondered whether this deficit is purely spatial, or it is influenced by the presence of a right-sided visual object. To answer this question, we tested 10 right brain-damaged patients with chronic left-neglect and 41 control participants on a cued response time (RT) detection task in which targets could appear in either of two lateral boxes. In different conditions, non-informative peripheral cues either consisted in the brightening of the contour of one lateral box (onset cue condition), or in the complete disappearance of one lateral box (offset cue condition). The target followed the cue at different stimulus-onset asynchronies (SOAs). If the disengagement deficit (DD) is purely space-based, then it should not vary across the two cueing conditions. With onset cues, patients showed a typical DD at short SOAs. With offset cues, however, the DD disappeared. Thus, patients did not show any DD when there was no object from which attention must be disengaged. These findings indicate that the attentional bias in left-neglect does not concern spatial locations per se, but visual objects in space.     

Covert Orienting of Visual Spatial Attention in Attention Deficit Hyperactivity Disorder: Does Comorbidity Make a Difference?

Attentional performance in children with attention deficit hyperactivity disorder (ADHD) with and without comorbid disorders was examined using the Covert Orienting of Visuospatial Attention Task (COVAT) and the Continuous Performance Task (CPT). The relationship between these two tasks was also examined. The results showed no overall differences on the attention tasks between children with ADHD alone and those with ADHD plus other disorders. Compared to non-ADHD control children, children with ADHD showed a deficit in the disengage operation of covert visuospatial attention, suggesting a difficulty in the endogenous mode of orienting. The ADHD children also showed a general performance deficit on the CPT. Although there was a general slowing on both attention tasks in the ADHD group, there was no relationship between invalid cue effect sizes on the COVAT and the CPT measures. These results indicate that these two attention tasks may be tapping both similar and independent underlying cognitive processes in ADHD.     

Processing Fragmented Forms and Strategic Control of Orienting in Visual Neglect

A single case study is reported of a patient with a right parietal lesion who showed: (1)clinical left neglect; (2)poor report of right-field items in identifying bilateral simultaneously presented stimuli; and (3)particularly poor processing of fragmented forms within his left visual field. We suggest that, in this patient, neglect is related to an impairment in orienting attention to contour information following his right-hemisphere lesion. This is exacerbated by the use of fragmented forms, but can be overcome by strategic allocation of attention to the left under bilateral presentation conditions. Neglect of the left, and extinction for right-side stimuli, reflect unbalanced competition on the attentional orienting system due respectively to (1) poor perceptual processing and (2) strategically attending to the left side of space.     

Saccadic Eye Movements in Object-based Neglect

RR showed left-sided neglect of objects following a stroke; when asked to copy a drawing of a simple scene he drew only the right side of each individual object. When his eye movements were recorded, RR made left saccades to locate objects to the left of the midline of the display, but restricted his fixations to the right sides of individual objects. Detailed examination of RR's eye movements while he viewed chimaeric photographs (comprising left and right halves of two different stimuli) also showed a pronounced tendency to scan only the right side of each. However, RR did not have a left hemianopia, and when left half stimuli were presented in isolation in his left visual field, he made left saccades and recognised them correctly. When presented with chimaeric stimuli aligned to the right of initial fixation, RR's first saccades landed on the left side of the images and a fixation of some 200m sec was made in the region of the left eye. Thus the initial left-sided fixation of chimaerics was unrelated to his ability to recognise them. Instead, on trials where left sides of chimaerics were recognised, RR made later saccades back into the left side of the chimaeric. In contrast, when left half stimuli were tachistoscopically presented at 200m sec exposure, RR could recognise a good proportion of them. We propose that RR has a deficit in orienting his attention within an object-based frame of reference, while being less impaired at orienting between objects. This does not result from a primary deficit of eye movement control; instead, RR's defective eye movements are a consequence of his object-based neglect.     

Ocular scanning and perceptual size distortion in hemispatial neglect: effects of prism adaptation and sequential stimulus presentation

When asked to compare two lateralized shapes for horizontal size, neglect patients often indicate the left stimulus to be smaller. Gainotti and Tiacci (1971) hypothesized that this phenomenon might be related to a rightward bias in the patients' gaze. This study aimed to assess the relation between this size underestimation and oculomotor asymmetries. Eye movements were recorded while three neglect patients judged the horizontal extent of two rectangles. Two experimental manipulations were performed to increase the likelihood of symmetrical scanning of the stimulus display. The first manipulation entailed a sequential, rather than simultaneous presentation of the two rectangles. The second required adaptation to rightward displacing prisms, which is known to reduce many manifestations of neglect. All patients consistently underestimated the left rectangle, but the pattern of verbal responses and eye movements suggested different underlying causes. These include a distortion of space perception without ocular asymmetry, a failure to view the full leftward extent of the left stimulus, and a high-level response bias. Sequential presentation of the rectangles and prism adaptation reduced ocular asymmetries without affecting size underestimation. Overall, the results suggest that leftward size underestimation in neglect can arise for a number of different reasons. Incomplete leftward scanning may perhaps be sufficient to induce perceptual size distortion, but it is not a necessary prerequisite.     

Responses to active and passive wrist rotation in area 5 of awake monkeys

In awake monkeys performing a wrist flexion-extension task, neurons which responded both dynamically and statically to joint rotation were found in contralateral area 5 of parietal cortex. Most received prominent input arising from deep receptors of the arm. The responses were directionally reciprocal and identical for passive and active joint rotations. In active movements, however, some firing patterns could not be fully explained in terms of sensory input alone. Thus, both peripheral and the postulated central inputs to area 5 may participate in the analysis of relative position of limb parts in space by this cortical region.     

DTI-MR tractography of white matter damage in stroke patients with neglect

Left visual neglect is a dramatic neurological condition that impairs awareness of left-sided events. Neglect has been classically reported after strokes in the territory of the right middle cerebral artery. However, the precise lesional correlates of neglect within this territory remain discussed. Recent evidence strongly suggests an implication of dysfunction of large-scale perisylvian networks in chronic neglect, but the quantitative relationships between neglect signs and damage to white matter (WM) tracts have never been explored. In this prospective study, we used diffusion tensor imaging (DTI) tractography in twelve patients with a vascular stroke in the right hemisphere. Six of these patients showed signs of neglect. Nonparametric voxel-based comparisons between neglect and controls on fractional anisotropy maps revealed clusters in the perisylvian WM and in the external capsule. Individual DTI tractography identified specific disconnections of the fronto-parietal and fronto-occipital pathways in the neglect group. Voxel-based correlation statistics highlighted correlations between patients' performance on two visual search tasks and damage to WM clusters. These clusters were located in the anterior limb of the internal capsule and in the WM underlying the inferior frontal gyrus, along the trajectory of the anterior segment of the arcuate fasciculus (asAF). These results indicate that chronic visual neglect can result from, and correlate with, damage to fronto-parietal connections in the right hemisphere, within large-scale cortical networks important for orienting of spatial attention, arousal and spatial working memory.     

Looking for the answer: the mind's eye in number space

Human subjects' answer to questions like "what number is halfway between 2 and 8" provides insights into spatial attention mechanisms involved in numerical processing. Here we show that mental numerical bisections are accompanied by a systematic pattern of horizontal eye movements: processing of a large number followed by a small number is accompanied with leftward eye movements, a tendency less pronounced or even reversed for the processing of a small number followed by a large number. The eyes thus appear to move along a left-to-right-oriented number line, indicating that shifts of attention in representational space are accompanied by an ocular motor orienting response. These results add to the growing evidence for a convergence of numerical processing, spatial attention, and movement planning in the parietal and frontal lobes. They also demonstrate the homologous relationship between our internal representations of numbers and space, and show that the concept of "number space" is more than a mere metaphor.     

Dissociation between physical and mental number line bisection in right hemisphere brain damage

To compare numeric quantities, humans make use of a 'mental number line' with smaller quantities located to the left of larger ones; it is unclear, however, whether orienting along the number line is like orienting along a physical line. We found that in brain-damaged subjects with defective leftward orienting, rightward deviation in the bisection of numeric and physical intervals is doubly dissociated. Deviation in numeric interval bisection was associated with prefrontal damage and spatial working memory deficit.     

Searching for the attention deficit in attention deficit hyperactivity disorder: the case of visuospatial orienting

We review all 14 extant studies of covert visuospatial attention in attention deficit hyperactivity disorder (ADHD) (total N=248). Metaanalysis showed that intriguing but isolated findings of alerting or posterior disengage deficits were too small to reliably detect with the sample sizes typically employed. Posterior move and engage operations and the vigilance sustained attention process were normal in ADHD. For exogenous cues, effect sizes for group differences were homogeneously small across all repeated-measures conditions, as were calculations of cost, benefit, and validity effects. For endogenous cues, effect sizes were heterogeneous; however, calculations of cost, benefit, and validity effects were small and homogenous. The most parsimonious conclusion may be that ADHD is not characterized by significant visual orienting dysfunction, but questions remain about the extent of anterior lateralized effects in the combined subtype and about attentional functioning in the inattentive subtype.     

Do patients with neglect show abnormal hand velocity profiles during tactile exploration of peripersonal space?

 It has been suggested that the movement impairments experienced by patients with neglect are not restricted to spatial disorders, but also affect higher-order kinematics (velocity and acceleration) to the extent that movements towards the neglected side are slower than movements away from it. In a recent study, we could not confirm this hypothesis, but found that patients with unilateral neglect exhibited no distinct direction-specific deficits in hand velocity when performing goal-directed reaching movements. Here we investigated whether neglect patients might reveal direction-specific deficits during exploratory hand movements. Six patients with left-sided neglect and six age-matched healthy control subjects scanned with their right hands the surface of a large table searching for a (non-existent) tactile target. Movements were performed in darkness. Time-position data of the hand were recorded with an optoelectronic camera system. Median activity of the patients’ exploratory hand movements was shifted to the right (Karnath and Perenin 1998). Hand trajectories were partitioned into sections of leftward/rightward or, along the sagittal plane, into sections of near/far movements. For each movement section average and peak velocities were computed. The patients’ hand movements were bradykinetic when compared with the control group. However, we found no evidence that average or peak velocities of leftward intervals were systematically lower than during rightward motion. Direction-specific deficits in velocity were also not observed for movements to and away from the body (sagittal plane). In conclusion, we found evidence for general bradykinesia in neglect patients but not for a direction-specific deficit in the control of hand velocity during exploratory hand movements. Received: 20 September 1998 / Accepted: 11 December 1998     

Oculographic analysis of word reading in hemispatial neglect

Neglect dyslexia is a disorder in which individuals misread text appearing on the contralateral side of space following an acquired lesion, usually to the right parietal lobe. This disorder is generally attributed to an impairment in representing spatial information. To determine whether the spatial representations underlying reading differ from those mediating other forms of visual behavior, we investigated the co-occurrence of neglect dyslexia with that of neglect, which manifests on tasks such as line bisection or line cancellation. We also examined the correlation between neglect dyslexia, when present, and eye movements in order to characterize the neglect dyslexia disorder further. Whereas there is no clear relationship between the reading disorder and other symptoms of visuospatial neglect, suggesting segregated spatial representations, there is a direct correspondence between the oculomotor performance of patients with neglect dyslexia and their reading behavior. This latter result suggests that the reading deficit may well arise from the failure to register and perceive the contralesional information.     

Phase-specific sensory representations in spinocerebellar activity during stepping: evidence for a hybrid kinematic/kinetic framework

The dorsal spinocerebellar tract (DSCT) provides a major mossy fiber input to the spinocerebellum, which plays a significant role in the control of posture and locomotion. Recent work from our laboratory has provided evidence that DSCT neurons encode a global representation of hindlimb mechanics during passive limb movements. The framework that most successfully accounts for passive DSCT behavior is kinematics-based having the coordinates of the limb axis, limb-axis length and orientation. Here we examined the responses of DSCT neurons in decerebrate cats as they walked on a moving treadmill and compared them with the responses passive step-like movements of the hindlimb produced manually. We found that DSCT responses to active locomotion were quantitatively different from the responses to kinematically similar passive limb movements on the treadmill. The differences could not be simply accounted for by the difference in limb-axis kinematics in the two conditions, nor could they be accounted for by new or different response components. Instead, differences could be attributed to an increased relative prominence of specific response components occurring during the stance phase of active stepping, which may reflect a difference in the behavior of the sensory receptors and/or of the DSCT circuitry during active stepping. We propose from these results that DSCT neurons encode two global aspects of limb mechanics that are also important in controlling locomotion at the spinal level, namely the orientation angle of the limb axis and limb loading. Although limb-axis length seemed to be an independent predictor of DSCT activity during passive limb movements, we argue that it is not independent of limb loading, which is likely to be proportional to limb length under passive conditions.     

Effects of unilateral frontal eye-field lesions on eye-head coordination in monkey

We studied the effects of unilateral frontal eye-field (FEF) lesions on eye-head coordination in monkeys that were trained to perform a visual search task. Eye and head movements were recorded with the scleral search coil technique using phase angle detection in a homogeneous electromagnetic field. In the visual search task all three animals showed a neglect for stimuli presented in the field contralateral to the lesion. In two animals the neglect disappeared within 2-3 wk. One animal had a lasting deficit. We found that FEF lesions that are restricted to area 8 cause only temporary deficits in eye and head movements. Up to a week after the lesion the animals had a strong preference to direct gaze and head to the side ipsilateral to the lesion. Animals tracked objects in contralateral space with combined eye and head movements, but failed to do this with the eyes alone. It was found that within a few days after the lesion, eye and head movements in the direction of the target were initiated, but they were inadequate and had long latencies. Within 1 wk latencies had regained preoperative values. Parallel with the recovery on the behavioral task, head movements became more prominent than before the lesion. Four weeks after the lesion, peak velocity of the head movement had increased by a factor of two, whereas the duration showed a twofold decrease compared with head movements before the lesion. No effects were seen on the duration and peak velocity of gaze. After the recovery on the behavioral task had stabilized, a relative neglect in the hemifield contralateral to the lesion could still be demonstrated by simultaneously presenting two stimuli in the left and right visual hemifields. The neglect is not due to a sensory deficit, but to a disorder of programming. The recovery from unilateral neglect after a FEF lesion is the result of a different orienting behavior, in which head movements become more important. It is concluded that the FEF plays an important role in the organization and coordination of eye and head movements and that lesions of this area result in subtle but permanent changes in eye-head coordination.     

The effects of competition and motor reprogramming on visuomotor selection in unilateral neglect

Patients with unilateral neglect following right hemisphere damage may have difficulty in moving towards contralesional targets. To test the hypothesis that this impairment arises from competing motor programs triggered by irrelevant ipsilesional stimuli, we examined 16 right hemisphere patients, eight with left visual neglect and eight without, in addition to eight healthy control subjects. In experiment 1 subjects performed sequences of movements using their right hand to targets on the contralesional or ipsilesional side of the responding limb. The locations of successive targets in each sequence were either predictable or unpredictable. In separate blocks of trials, targets appeared either alone or with a simultaneous distractor located at the immediately preceding target location. Neglect patients were significantly slower to execute movements to contralesional targets, but only for unpredictable movements and in the presence of a concurrent ipsilesional distractor. In contrast, healthy controls and right hemisphere patients without neglect showed no directional asymmetries of movement execution. In experiment 2 subjects were required to interrupt a predictable, reciprocating sequence of leftward and rightward movements in order to move to an occasional, unpredictable target that occurred either in the direction opposite to that expected, or in the same direction but twice the extent. Neglect patients were significantly slower in reprogramming the direction and extent of movements towards contralesional versus ipsilesional targets, and they also made significantly more errors when executing such movements. Right hemisphere patients without neglect showed a similar bias in reprogramming direction (but not extent) for contralesional targets, whereas healthy controls showed no directional asymmetry in either condition. On the basis of these findings we propose that neglect involves a competitive bias in favour of motor programs for actions directed towards ipsilesional versus contralesional events. We suggest that programming errors and increased latencies for contralesional movements arise because the damaged right hemisphere can no longer effectively inhibit the release of inappropriate motor programs towards ipsilesional events. Received: 1 October 1996 / Accepted: 21 October 1997     

Brainstem control of head movements during orienting; organization of the premotor circuits

When an object appears in the visual field, animals orient their head, eyes, and body toward it in a well-coordinated manner (orienting movement). The head movement is a major portion of the orienting movement. Interest in the neural control of head movements in the monkey and human have increased in the 1990's, however, fundamental knowledge about the neural circuits controlling the orienting head movement continues to be based on a large number of experimental studies performed in the cat. Thus, it is crucial now to summarize information that has been clarified in the cat for further advancement in understanding the neural control of head movements in different animal species. The superior colliculus (SC) has been identified as the primary brainstem center controlling the orienting. Its output signal is transmitted to neck motoneurons via two major separate pathways: one through the reticulospinal neurons (RSNs) in the pons and medulla and the other through neurons in Forel's field H (FFH) in the mesodiencephalic junction. The tecto-reticulo-spinal pathway controls orienting chiefly in the horizontal direction, while the tecto-FFH-spinal pathway controls orienting in the vertical direction. In each pathway, a subgroup of neurons functions as premotor neurons for both extraocular and neck motoneurons, while others are specified for each, which allows both coordinated and separate control of eye and head movements. Head movements almost always produce shifts in the center of gravity that might cause postural disturbances. The postural equilibrium may be maintained by transmitting the orienting command to the limb segments via descending axons of the reticulospinal and long propriospinal neurons. The SC and brainstem relay neurons receive descending inputs from higher order structures such as the cerebral cortex, cerebellum, and basal ganglia. These inputs may serve context-dependent control of orienting by modulating the activities of the primary brainstem pathways.     

Dissociation between visual line bisection and mental number line bisection in schizophrenia

Many studies of hemispatial neglect patients have indicated that spatial attention processes operate similarly in visual space and number space. However, some studies have indicated a dissociation of processing between visual line bisection and mental number bisection. A number of investigations have suggested that schizophrenic patients show a mild right pseudo-neglect on visual line bisection tasks. The present study was designed to determine if a functional link exists between performance of visual line and number line bisection in schizophrenic patients. Groups of 40 schizophrenic patients and 40 controls performed each bisection task. In the visual line bisection task, schizophrenic patients showed a significant leftward bias relative to the healthy controls for 9 different line lengths. No significant difference in bias was found between the 2 groups on the mental line bisection task. These results indicated that schizophrenic patients may exhibit attention deficit with respect to visual space but not number space, suggestive of the dissociation of processing between visual line bisection and mental number line bisection. These results provide more insight into the correlation between the visual line and number bisection tasks in schizophrenic patients.     

A lesion analysis of visual orienting performance in children with cerebral vascular injury

Anterior brain insults during development have been shown to result in visual orienting deficits; however, the type of orienting deficit has varied across studies. Performance on an orienting task was examined in relation to the location and volume of injury on magnetic resonance exams in 15 children with cerebral infarction and 32 control children. Contralateral lesions including the parietal lobe were associated with larger validity effects, suggesting difficulties disengaging attention. A similar trend was found for the middle-frontal gyrus. Basal ganglia injury contralateral to a given hemifield was associated with less facilitation of attention. Lesion volume in each hemisphere did not show a significant relation with contralateral validity effects. The data suggest that variability in the type of visual orienting deficits shown in prior studies of children with anterior brain insults may be related in part to the specific location of lesions within the frontal lobe.