Spatial orientation and the representation of space with parietal lobe lesions

Damage to the human parietal cortex leads to disturbances of spatial perception and of motor behaviour. Within the parietal lobe, lesions of the superior and of the inferior lobule induce quite different, characteristic deficits. Patients with inferior (predominantly right) parietal lobe lesions fail to explore the contralesional part of space by eye or limb movements (spatial neglect). In contrast, superior parietal lobe lesions lead to specific impairments of goal-directed movements (optic ataxia). The observations reported in this paper support the view of dissociated functions represented in the inferior and the superior lobule of the human parietal cortex. They suggest that a spatial reference frame for exploratory behaviour is disturbed in patients with neglect. Data from these patients' visual search argue that their failure to explore the contralesional side is due to a disturbed input transformation leading to a deviation of egocentric space representation to the ipsilesional side. Data further show that this deviation follows a rotation around the earth-vertical body axis to the ipsilesional side rather than a translation towards that side. The results are in clear contrast to explanations that assume a lateral gradient ranging from a minimum of exploration in the extreme contralesional to a maximum in the extreme ipsilesional hemispace. Moreover, the failure to orient towards and to explore the contralesional part of space appears to be distinct from those deficits observed once an object of interest has been located and releases reaching. Although patients with neglect exhibit a severe bias of exploratory movements, their hand trajectories to targets in peripersonal space may follow a straight path. This result suggests that (i) exploratory and (ii) goal-directed behaviour in space do not share the same neural control mechanisms. Neural representation of space in the inferior parietal lobule seems to serve as a matrix for spatial exploration and for orienting in space but not for visuomotor processes involved in reaching for objects. Disturbances of such processes rather appear to be prominent in patients with more superior parietal lobe lesions and optic ataxia.     

Spiroperidol reinstates asymmetries in neglect in rats recovered from left or right dorsomedial prefrontal cortex lesions.

The role of dopaminergic mechanisms in spontaneous behavioral recovery from cortical neglect was examined in rats that received lesions of either the left or the right dorsomedial prefrontal cortex (PCm, also referred to as AGm). Neglect was assessed by rating the degree of head orientation to visual, auditory, or tactile stimuli. Following behavioral recovery, separate groups received 0.03, 0.05, 0.07, or 0.10 mg/kg spiroperidol or the vehicle. In accordance with the lateralization of neglect typically seen postsurgery, spiroperidol dose-dependently reinstated contralesional neglect in left PCm operates and ipsilesional neglect in right PCm operates. Neglect was reinstated by spiroperidol in right PCm operates at lower doses than in left PCm operates. Also, only right PCm operates demonstrated asymmetrical bilateral dose-dependent neglect. Spiroperidol did not produce neglect in unilaterally brain-damaged control subjects. The results indicate that dopaminergic mechanisms may underlie spontaneous recovery from cortical neglect and that these mechanisms are asymmetrical in left and right PCm operates. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Independent practice association physician groups in California

Hemispatial neglect is a neurological disorder which entails a spatial bias that penalizes events occurring in the hemispace contralateral to a brain lesion. Mechanisms operating upon various stages ranging from perception to action have been invoked to explain neglect. The present study explores the contribution of a defective programming of arm movements towards the neglected hemispace to neglect behaviour. Two reaction time tasks -- a "perceptual" task and a "motor" task -- were performed by right brain-damaged (RBD) patients with left hemispatial neglect, RBD patients without signs of neglect and control subjects. The perceptual task consisted of lateralized visual stimuli and central motor responses, whereas the motor task consisted of visual stimuli presented on the vertical midline and hand responses to be produced in either hemispace. Neglect patients showed a rightward bias on the perceptual task, but only two RBD patients (showing no signs of severe neglect) were consistently slowed in producing leftward motor responses. Different reference frames may thus be used in perceptual tasks and tasks involving arm movements. We conclude that hemispatial neglect commonly results from attentional impairments operating upon a visual perceptual frame of reference; additional deficits appear to be necessary to produce a directional motor disorder.     

Response channel activation and the temporoparietal junction

When we learn to make one motor response to one visual stimulus and a different motor response to another, representations of these stimulus-response associations must be maintained to efficiently transduce perception into action. When an irrelevant distractor is presented adjacent to a target stimulus, interference is observed when the two stimuli are associated with conflicting responses, presumably due to response channel activation by the incompatible information. We have explored the neural bases of these interference effects. In a previous study, patients with hemispatial neglect showed normal interference from contralesional flankers. In another study, patients with lesions of the lateral prefrontal cortex were found not to show interference from distractors presented in the contralesional hemifield. The current study provided a more anatomically detailed investigation of the effects of posterior association cortex lesions on flanker interference. Patients with chronic, unilateral lesions involving the temporoparietal junction (TPJ), two of whom had hemispatial neglect, were compared with patients with lesions of the posterior association cortex not involving the TPJ. All patients performed a color discrimination task at fixation while a congruent or incongruent colored flanker was briefly presented (16.7 ms) in the adjacent contralesional or ipsilesional hemifield. Patients with TPJ lesions showed no interference effects from the contralesional flankers. These results suggest that the TPJ, in combination with the dorsolateral prefrontal cortex, is involved in transducing perception into action.     

Patterns of dissociation between left hemineglect and deviation of the egocentric reference

Sixteen control subjects and six right brain-damaged patients with left hemiparesis (three showing signs of left unilateral neglect, three with no signs of neglect) performed a straight-ahead pointing task with their right hand while blindfolded. The aim was to test the hypothesis that the egocentric reference shows significant ipsilesional deviation in left neglect patients. We found no correlation between the position of the egocentric reference and the presence of neglect signs. Neglect patients, like non-neglect patients, showed leftward, rightward or no significant deviation when pointing straight ahead. Results are discussed with reference to egocentric hypotheses of neglect and experimental remission of neglect.     

Seeing only the right half of the forest but cutting down all the trees?

Unilateral neglect following damage to the right hemisphere of the brain can be characterized by failure of the global attentional mechanisms of the right hemisphere to direct the local detail processors of the left hemisphere towards the contralesional left hemispace. This is suggested by patients who recognize the global form of the left side of shapes (the forest) but fail to cancel out its local details (the trees). Here we report the opposite behavioural dissociation in a patient (Q.M.) with damage to the right hemisphere of the brain. Q.M. detected local details (such as the tail of a dog) on the left or right side of visual shapes, regardless of whether these details belonged to predefined target shapes (a dog in this case) or to distractor shapes differing on the opposite side (a dog with a swan's neck and head, for example). Psychological testing showed an abnormal tendency of this patient to respond to local features, but perfect accuracy in interpreting global features when the local features could not interfere in global processing. The results indicate that the left hemisphere can integrate multiple local features simultaneously but loses global awareness as soon as local features individually compete for response selection. However, awareness of the whole is not necessary for the sequential processing of the parts.     

Preattentive and attentive visual search in individuals with hemispatial neglect.

Preattentive and attentive visual processing was examined in patients with hemispatial neglect, hemispatial neglect with hemianopia, and control participants. In the preattentive search task, targets possessed a unique feature that was not shared by distractors. In the attentive search task, targets lacked a feature that was present in the distractors. Preattentive search was normal in 3 neglect patients with cortical lesions but not in 2 neglect patients with hemianopia. A 4th neglect patient without hemianopia with a subcortical infarct abnormally used serial search mechanisms in the preattentive task. Neglect patients were characteristically impaired in the contralesional field in the attentive search task. This study demonstrates preserved explicit detection of visual features in cases of hemispatial neglect. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Search deficits in neglect patients are dependent on size of the visual scene.

Patients with hemispatial neglect are impaired at moving their attention to a target on the contralesional display side. In the present study, visual display area was varied independently of the number of items displayed within the area. Patients searched for the absence of a simple visual feature in displays that produce serial search performance in normals. The contralesional delay was enhanced for stimulus arrays that were distributed over a larger display area, which suggests that neglect is more severe when attention has to be moved over a larger distance. The directional disengage deficit in neglect therefore depends on the target's relative position within a display, the number of ipsilesional items competing for attention, and the distance of the contralesional attention movements. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Hand, space and attentional asymmetries in goal-directed manual aiming

Two experiments were conducted to explore the interaction of the two cerebral hemispheres in motor control, by examining hand, space and attentional asymmetries in goal-directed aiming. In Experiment 1, right-handed subjects moved to targets more quickly with their right hand than their left hand. In addition, each hand was faster when moving in its own hemispace. Although in a control condition, movements were initiated more quickly with the left hand, visual distractors disrupted left hand performance more than right hand performance. For contralateral aiming, ipsilateral distractors caused the greatest interference. In Experiment 2, when targets and distractors were all presented at the midline, a right hand advantage was found for movement time along with a left hand advantage for reaction time, independent of target and distractor location. Our findings are discussed in terms of a right hemisphere role in movement preparation and the allocation of attention in space, and greater left hemisphere involvement in movement execution.     

Disentangling gravitational, environmental, and egocentric reference frames in spatial neglect

Previous studies in neglect patients using rotation of the body around the roll-axis revealed neglect of visual stimuli not only in the egocentric, body-centered left but also in the environmental left. The latter has been taken as evidence for a gravity-based environment-centered component of neglect occurring independently of the subject's actual body orientation. However, by using visual stimuli in a normally lightened room, the studies confounded the gravitational upright with the visible upright of the surround. Thus, it is possible that the visible upright of the environment may have served the role of the gravitational upright relative to which neglect occurred. The present experiment evaluated the influence of gravity on contralateral neglect when no visual information was presented. In complete darkness, neglect patients' exploratory eye movements were recorded in five experimental conditions: body in normal upright position, body titled 30 degrees to the left and 30 degrees to the right, and body pitched 30 degrees backward and 30 degrees forward. In the upright orientation, the patients with neglect showed a bias of ocular exploration to the ipsilesional right side. In egocentric body coordinates, we found no significant differences between the orientation of the biased search field in the different experimental conditions showing that the search field shifted with the orientation of the body. No significant decrease or enhancement of neglect was observed when body orientation was varied in the different conditions. In conclusion, the present results revealed that the modulation of gravitational forces has no significant influence on the exploratory bias of these patients. When visual information was excluded and only graviceptive information was available, the patients' failure to explore the contralesional part of space appeared purely body-centered. The results argue against a disturbed representation of space in neglect that encodes locations in a gravity-based reference system.     

Reaching to ipsilateral or contralateral targets: within-hemisphere visuomotor processing cannot explain hemispatial differences in motor control

Aiming movements made to visual targets on the same side of the body as the reaching hand typically show advantages as compared to aiming movements made to targets on the opposite side of the body midline in the contralateral visual field. These advantages for ipsilateral reaches include shorter reaction time, higher peak velocity, shorter duration and greater endpoint accuracy. It is commonly hypothesized that such advantages are related to the efficiency of intrahemispheric processing, since, for example, a left-sided target would be initially processed in the visual cortex of the right hemisphere and that same hemisphere controls the motor output to the left hand. We tested this hypothesis by examining the kinematics of aiming movements made by 26 right-handed subjects to visual targets briefly presented in either the left or the right visual field. In one block of trials, the subjects aimed their finger directly towards the target; in the other block, subjects were required to aim their movement to the mirror symmetrical position on the opposite side of the fixation light from the target. For the three kinematic measures in which hemispatial differences were obtained (peak velocity, duration and percentage of movement time spent in deceleration), the advantages were related to the side to which the motor response was directed and not to the side where the target was presented. In addition, these effects tended to be larger in the right hand than in the left, particularly for the percentage of the movement time spent in deceleration. The results are interpreted in terms of models of biomechanical constraints on contralateral movements, which are independent of the hemispace of target presentation.     

Deficits and recovery of first- and second-order motion perception in patients with unilateral cortical lesions

Unilateral lesions in the posterior parietal cortex can degrade motion perception in the contralesional visual hemifield. Our aim was to investigate whether deficits caused by cortical lesions may be different for first- and second-order motion perception, and to study the time scale of any potential recovery. In nine patients with circumscribed lesions mainly in the parietal and fronto-parietal cortex, thresholds for direction discrimination were measured for stimuli presented peripherally in their ipsi- and contralesional hemifield. Subjects had to identify the direction of a vertically moving object embedded in a background of dynamic random dot noise. The object consisted of various proportions of signal and noise dots. Signal dots were either (a) coherently moving in the same direction as the object (first-order), (b) stationary (second-order: drift-balanced), or (c) coherently moving in the opposite direction (second-order: theta). Noise dots were flickering. Two patients showed significant threshold elevations for all three types of motion stimuli presented in their contralesional hemifield, while thresholds for ipsilesional targets were unaffected. Neither showed any selective deficit of first- versus second-order motion perception, but second-order motion was more impaired. Their lesions probably included the motion area V5-MT, which was spared in the other seven patients. One of the patients, who was retested several times during a 27-month postlesional period, showed complete recovery for first- and second-order motion direction discrimination, as well as for the detection of speed differences.     

Dopamine agonists reorient visual exploration away from the neglected hemispace

OBJECTIVE: To study the effects of bromocriptine, a dopamine agonist, on visual search. BACKGROUND: The anatomic substrate of spatial attention takes the form of a distributed network with interconnected cortical (frontal, parietal, and cingulate) and subcortical (striatal and thalamic) components. Dopamine appears to exert a modulatory effect on the function of this network. METHODS: Seven consecutive right-handed subjects with right-sided cerebral lesions were studied using a computerized target search paradigm. Eye movements were recorded. RESULTS: Bromocriptine caused the subjects to spend more time exploring the ipsilesional hemispace and therefore increased the relative neglect of the contralesional left hemispace. However, target detection accuracy did not change. Bromocriptine thus had a differential impact on the exploratory-motor versus sensory-perceptual components of directed attention. CONCLUSIONS: Our results show that bromocriptine may worsen some aspects of hemispatial neglect in patients with lesions that include the postsynaptic components of ascending dopaminergic pathways.     

Time perception in spatial neglect: A distorted representation?

Objective: It has been proposed that time, space, and numbers share the same metrics and cortical network, the right parietal cortex. Several recent investigations have demonstrated that the mental number line representation is distorted in neglect patients. The aim of this study is to investigate the relationship between time and spatial configuration in neglect patients. Method: Fourteen right-brain damaged patients (six with neglect and eight without neglect), as well as eight age-matched healthy controls, performed a time discrimination task. A standard tone (short: 700 ms and long: 1,700 ms) had to be confronted in duration to a test tone. Test tone differed of 100, 200, and 300 ms respect to the standard tone duration. Results: Neglect patients performed significantly worse than patients without neglect and healthy controls, irrespective of the duration of the standard tone. Conclusion: These results support the hypothesis that mental representations of space and time both share, to some extent, a common cortical network. Besides, spatial neglect seems to distort the time representation, inducing an overestimation of time durations. (PsycINFO Database Record (c) 2011 APA, all rights reserved)     

Shifting visual attention between objects and locations: Evidence from normal and parietal lesion subjects.

Space- and object-based attention components were examined in neurologically normal and parietal-lesion Ss, who detected a luminance change at 1 of 4 ends of 2 outline rectangles. One rectangle end was precued (75% valid); on invalid-cue trials, the target appeared at the other end of the cued rectangle or at 1 end of the uncued rectangle. For normals, the cost for invalid cues was greater for targets in the uncued rectangle, indicating an object-based component. Both right- and left-hemisphere patients showed costs that were greater for contralesional targets. For right-hemisphere patients, the object cost was equivalent for contralesional and ipsilesional targets, indicating a spatial deficit, whereas the object cost for left-hemisphere patients was larger for contralesional targets, indicating an object deficit. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Apoptosis signaling by death receptors

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.     

In search of biased egocentric reference frames in neglect

The present study was aimed at assessing, by means of visual as well as proprioceptive-kinaesthetic straight-ahead tasks, the possible causal role of the ipsilesional deviation of the egocentric reference frame in determining neglect syndrome. The hypothesis, originally proposed by Ventre et al. [3], that an alteration of the representation of body-centred space can be a cause of asymmetrical spatial behaviour in humans has been recently revived by Karnath and co-workers [24]. The results of the present study seem to challenge the view that a systematic ipsilesional displacement of the egocentric reference is the crucial mechanism responsible for unilateral visual neglect. Under visual conditions, in which patients were required to stop a moving spot as it crossed their perceived midline, the ipsilesional deviation of the egocentric reference frame was dependent upon the direction of visual scanning. Right to left visual scanning direction produced a rightward displacement of the egocentric reference. In contrast, left to right visual scanning direction allowed neglect patients to correctly locate their perceived egocentre with an accuracy which did not differ from controls. The notion that the effect of a deviation of the egocentric reference frame is actually dependent on a bias in the visual scanning orienting response was also confirmed in the proprioceptive straight-ahead pointing tasks, in which the patients were blindfolded and therefore no visual information was available. In these conditions, in which patients were required to judge the subjective midline by using head, trunk and shoulder co-ordinate systems, the displacement of the subjective egocentric midline was not present.     

Visuomotor processing in unilateral neglect

The extent to which visual information on the contralateral, unattended side influences the performance of patients with hemispatial neglect was studied in a visuomotor reaching task. We replicated the well-established finding that, relative to target-alone trials, normal subjects are slower to reach to targets in the presence of visual distractors which appear either ipsilateral or contralateral to the target, with greater interference in the former condition. Six patients with hemispatial neglect showed even greater interference than did the normal subjects when the distractor appeared ipsilaterally but showed no significant interference from contralateral distractors. This pattern of performance was qualitatively similar for patients with lesions restricted to posterior regions and for patients with more extensive lesions involving both posterior and anterior brain regions. These findings suggest that, in the visuomotor domain, information on the contralateral side is processed minimally, if at all, in patients with hemispatial neglect.     

Object-based inhibition of return in patients with posterior parietal damage.

Recent evidence has shown that inhibition of return, IOR, is impaired in patients with parietal damage with or without clinical signs of neglect (Bartolomeo, Sieroff, Decaix, & Chokron, 2001; Vivas, Humphreys, & Fuentes, 2003, respectively). In addition to environment-based IOR, Tipper et al. (1991) showed that IOR could be also associated with dynamic, object-based representations. In our study, we examined four patients with unilateral lesions to the parietal lobe, and a group of healthy controls, in an IOR procedure with moving objects where a pre-cued object could move, clockwise or counterclockwise, 90° in polar coordinates. The group of control participants showed a small but significant object-based IOR effect. In contrast, the patients showed an object-based IOR effect when the objects moved from the contralesional field toward the ipsilesional field, whereas there was no IOR effect when they moved from the ipsilesional to the contralesional field. These findings are discussed in terms of the role of the parietal cortex in implementing attentional biases in both environment-based (Vivas et al., 2003) and object-based frames of reference. (PsycINFO Database Record (c) 2010 APA, all rights reserved)     

Saccadic eye movement and working memory deficits following damage to human prefrontal cortex

A patient with a lesion confined largely to the right inferior frontal gyrus was found to be impaired on tests of spatial working memory and executive functioning. By contrast, his pattern recognition was good. The patient's selective impairments are consistent with the view that prefrontal cortex contributes to processes involved in spatial working memory. The patient was also tested on a range of oculomotor paradigms, some of which required the temporary suppression of a saccadic response. He was unable to suppress making contra- or ipsilesional reflexive glances to peripheral stimuli on the "anti-saccade" paradigm, but his performance improved on delayed saccade, memory-guided saccade and fixation tasks. Although reflexive glances were observed under these conditions they occurred more frequently in response to contralesional stimuli than ipsilesional ones. Furthermore, the patient had no difficulty in performing anti-point movements with his ipsilesional hand. Thus, his inability to suppress reflexive glances on the anti-saccade task is not due to a generalised problem of "distractibility". The patient's deficits are discussed in terms of models of anti-saccade generation and are related to recent findings regarding the role of prefrontal cortex in working memory and visual attention.