Correlated deficits of perception and action in optic ataxia.

Optic ataxia, following dorsal stream lesions, is characterised by impaired visuomotor guidance. Recent studies have found concurrent perceptual deficits, but it is unclear whether these are functionally related to the visuomotor symptoms. We studied the ability of a well-documented patient (IG) with bilateral optic ataxia to react to sudden target jumps by correcting ongoing reaches or by explicitly reporting the jump direction. IG showed deficient reach corrections, especially for target jumps to the visual periphery, and was similarly slow to discriminate the same jumps perceptually. Across six test conditions, in which the retinal locations of target jumps were varied, her perceptual slowing mirrored her reaching deficit precisely. These findings confirm perceptual impairments after dorsal stream lesions, and imply a shared functional basis with the classical visuomotor symptoms of optic ataxia. Additionally, we show that the online correction deficit is determined dually by the retinal location to which the reach must be diverted, and the location to which it is initially directed. We suggest that this deficit, and its perceptual counterpart, can be traced to a slowed contralesional orienting of attention in optic ataxia.

Choosing between alternative wrist postures: action planning needs perception.

When normal subjects grasp with their right hand a rectangular object placed at different orientations in the horizontal plane, they change from a 'thumb left' (clockwise) to a 'thumb right' (anti-clockwise) grasp when the orientation exceeds about 110 degrees , with respect to the mid-sagittal plane. This suggests planning of the final grip orientation at, or before the start of the prehension movement. The current study assessed performance of two visual agnosic patients (SB and DF) on a grasping task requiring the planning of final grip posture. Five healthy subjects were also tested. Subjects were required to grasp a triangular-section block, which was presented at one of seven different orientations (80-140 degrees). The healthy subjects showed a consistent relation between object orientation and hand orientation just before contact. In addition, they consistently used a clockwise grasp when object orientation was less than 100 degrees , and an anti-clockwise grasp when it was more than 110 degrees, with a sharply defined switch-point being identifiable for each subject. For both visual agnosic patients, hand orientation was also reliably related to object orientation. However, the selection of grasp posture was markedly abnormal: they did not consistently switch between clockwise and anti-clockwise grasps within the normal orientation range, and the switch, when it did occur, was not at all sharply defined. These results suggest that the planning of hand orientation during a grasp depends on a perceptually based judgement of the awkwardness of alternative movements. This would presumably involve ventral stream processing, which is disrupted in the visual agnosic patients.

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.

Reaching errors in optic ataxia are linked to eye position rather than head or body position.

When reaching towards a visual stimulus, spatial information about the target must be transformed into an appropriate motor command. Visual information is coded initially in retinotopic coordinates, while the reaching movement ultimately requires the specification of the target position in limb-centred coordinates. It is well established that the posterior parietal cortex (PPC) plays an important role in transforming visual target information into motor commands. Lesions in the PPC can result in optic ataxia, a condition in which the visual guidance of goal-directed movements is impaired. Here, we present evidence from two patients with unilateral optic ataxia following right PPC lesions, that the pattern of reaching errors is linked to an eye-centred frame of reference. Both patients made large errors when reaching to visual targets on the left side of space, while facing and fixating straight ahead. By varying the location of fixation and the orientation of the head and body, we were able to establish that these large errors were made specifically to targets to the left of eye-fixation, rather than to the left of head-, body-, or limb-relative space. These data support the idea that visual targets for reaching movements are coded in eye-centred coordinates within the posterior parietal cortex.

Sensorimotor effects on central space representation: prism adaptation influences haptic and visual representations in normal subjects.

Prism adaptation improves visual and haptic manifestations of left neglect, and can induce a small but reliable simulation of left visual neglect in normal individuals. Here, we present two experiments in which the effects of prism adaptation on the representation of space were explored. In Experiment 1, normal subjects were required to locate the centre of a haptically explored circle, before and after adaptation to leftward displacing prisms. In Experiment 2, a visual circle centring task was used. In both tasks, prism adaptation induced a significant rightward shift of performance. In addition, in both experiments, three classical measures of visuo-manual adaptation were taken: the visual shift, the proprioceptive shift and the total shift. The effects found on the haptic and visual tasks did not correlate with any of these measures. This suggests that the effects of prism adaptation on the circle centring tasks did not depend directly on the sensorimotor consequences of the adaptation. These results imply that prism adaptation can affect noetic levels of space representation in normal subjects, supporting the hypothesis that this low-level sensorimotor intervention can exert a bottom-up structuring influence on higher levels of cognitive integration.

Preserved obstacle avoidance during reaching in patients with left visual neglect.

We asked 12 patients with left visual neglect to bisect the gap between two cylinders or to reach rapidly between them to a more distal target zone. Both tasks demanded a motor response but these responses were quite different in nature. The bisection response was a communicative act whereby the patient indicated the perceived midpoint. The reaching task carried no imperative to bisect the gap, only to maintain a safe distance from either cylinder while steering to the target zone. Optimal performance on either task could only be achieved by reference to the location of both cylinders. Our analysis focused upon the relative influence of the left and right cylinders on the lateral location of the response. In the bisection task, all neglect patients showed qualitatively the same asymmetry, with the left cylinder exerting less influence than the right. In the reaching task, the neglect group behaved like normal subjects, being influenced approximately equally by the two cylinders. This was true for all bar two of the patients, who showed clear neglect in both tasks. We conclude that the visuomotor processing underlying obstacle avoidance during reaching is preserved in most patients with left visual neglect.

Avoidance of obstacles in the absence of visual awareness.

The spatial character of our reaching movements is extremely sensitive to potential obstacles in the workspace. We recently found that this sensitivity was retained by most patients with left visual neglect when reaching between two objects, despite the fact that they tended to ignore the leftward object when asked to bisect the space between them. This raises the possibility that obstacle avoidance does not require a conscious awareness of the obstacle avoided. We have now tested this hypothesis in a patient with visual extinction following right temporoparietal damage. Extinction is an attentional disorder in which patients fail to report stimuli on the side of space opposite a brain lesion under conditions of bilateral stimulation. Our patient avoided obstacles during reaching, to exactly the same degree, regardless of whether he was able to report their presence. This implicit processing of object location, which may depend on spared superior parietal-lobe pathways, demonstrates that conscious awareness is not necessary for normal obstacle avoidance.

Delayed reaching and grasping in patients with optic ataxia.

A series of experiments documenting the reaching and grasping of two patients with optic ataxia is presented. We compare their immediate responses with their behavior when required to delay for a few seconds before responding. When the delayed response is 'pantomimed', i.e. made in the absence of the target object, their performance typically improves. This pattern was predicted from a two-visual-systems model in which the cortical dorsal stream mediates normal visually guided actions while the ventral stream deals with visual information that has to be held in memory. We further found that when a 'preview' task was used in which the patients could use memorized information to guide a response to a still-present target object, they did so in preference to using the visual information facing them.

Disentangling perceptual and response bias in unilateral neglect: a methodological proposal.

Bisiach and his colleagues have developed a new version of Milner's "landmark task" for the purpose of separating "perceptual" and "response" biases in neglect patients. Subjects are required to decide which is the longer (or the shorter) of the two portions of a pre-bisected horizontal line. The authors proposed two indices to measure perceptual and response bias respectively. However, these indices are not mathematically independent of one another. Moreover, they do not exploit all of the information available in the data, since they do not consider the effect of the different transection locations across trials. We now propose an alternative means of analyzing data from the revised Landmark task, which generates independent estimates of perceptual and response biases. The method and its theoretical foundation are summarized, and illustrative data obtained from brain damaged patients and control subjects are presented.

Visual and tactile size distortion in a patient with right neglect.

One typical feature of the neglect syndrome in patients with right hemisphere damage is that they bisect horizontal lines to the right of centre. It has been argued that to a large extent these bisection errors can be attributed to a perceptual change whereby the patient experiences the left half of a line as shorter than the right half, causing them to set the midpoint of the line towards the right. We describe here a patient with a left hemisphere lesion and rightward neglect, who consequently makes bisection errors in a leftward direction. We carried out a series of tests which confirmed that he shows a subjective visual distortion in the converse direction, i.e. a perception of horizontal extents on the right as shorter than extents on the left. We also found that he shows a similar distortion in his tactile perception. The association of visual and tactile distortions in this patient is compatible with the view that the distortion effects have a rather high-level origin. Multiple single-case studies will, however, be required to establish whether this association of deficits is typical, or whether visual and tactile size distortions are separable symptoms associated with neglect.

Grasping the past. delay can improve visuomotor performance.

"Optic ataxia" is caused by damage to the human posterior parietal cortex (PPC). It disrupts all components of a visually guided prehension movement, not only the transport of the hand toward an object's location, but also the in-flight finger movements pretailored to the metric properties of the object. Like previous cases, our patient (I.G.) was quite unable to open her handgrip appropriately when directly reaching out to pick up objects of different sizes. When first tested, she failed to do this even when she had previewed the target object 5 s earlier. Yet despite this deficit in "real" grasping, we found, counterintuitively, that I.G. showed good grip scaling when "pantomiming" a grasp for an object seen earlier but no longer present. We then found that, after practice, I.G. became able to scale her handgrip when grasping a real target object that she had previewed earlier. By interposing catch trials in which a different object was covertly substituted for the original object during the delay between preview and grasp, we found that I.G. was now using memorized visual information to calibrate her real grasping movements. These results provide new evidence that "off-line" visuomotor guidance can be provided by networks independent of the PPC.

Monocular and binocular distance cues: insights from visual form agnosia I (of III).

The human nervous system constructs a Euclidean representation of near (personal) space by combining multiple sources of information (cues). We investigated the cues used for the representation of personal space in a patient with visual form agnosia (DF). Our results indicated that DF relies predominantly on binocular vergence information when determining the distance of a target despite the presence of other (retinal) cues. Notably, DF was able to construct an Euclidean representation of personal space from vergence alone. This finding supports previous assertions that vergence provides the nervous system with veridical information for the construction of personal space. The results from the current study, together with those of others, suggest that: (i) the ventral stream is responsible for extracting depth and distance information from "monocular" retinal cues (i.e. from shading, texture, perspective) and (ii) the dorsal stream has access to binocular information (from horizontal image disparities and vergence). These results also indicate that DF was not able to use size information to gauge target distance, suggesting that intact temporal cortex is necessary for "learned size" to influence distance processing. Our findings further suggest that in neurologically intact humans, object information extracted in the ventral pathway is combined with the products of dorsal stream processing for guiding prehension. Finally, we studied the "size-distance paradox" in visual form agnosia in order to explore the cognitive use of size information. The results of this experiment were consistent with a previous suggestion that the paradox is a cognitive phenomenon.

The role of size and binocular information in guiding reaching: insights from virtual reality and visual form agnosia III (of III).

Reaching out to grasp an object requires information about the size of the object and the distance between the object and the body. We used a virtual reality system with a control population and a patient with visual form agnosia (DF) in order to explore the use of binocular information and size cues in prehension. The experiments consisted of a perceptual matching task in addition to a prehension task. In the prehension task, control participants modified their reach distance in response to step changes in vergence in the absence of any clear reference for relative disparity. Their reach distance was unaffected by equivalent step changes in size, even though they used this information to modify grasp and showed a size bias in a distance matching task. Notably, DF showed the same pattern of results as the controls but was far more sensitive to step changes in vergence. This finding complements previous research suggesting that DF relies predominantly on vergence information when gauging target distance. The results from the perceptual matching tasks confirmed previous findings suggesting that DF is unable to make use of size information for perceptual matching, including distance comparisons. These data are discussed with regard to the properties of the pathways subserving the two visual cortical processing streams.

Vertical gaze angle as a distance cue for programming reaching: insights from visual form agnosia II (of III).

It has been shown that a patient with visual form agnosia (DF) relies predominantly on vergence information when gauging target distance in an open-loop pointing task. This finding suggested that the programming of prehension might be severely disrupted if DF viewed target objects through ophthalmic prisms. An initial experiment showed that this prediction was not upheld; DF was able to programme reasonably accurate movements to objects located on a tabletop despite large changes in vergence angle. A second experiment, however, showed that placing the target objects at eye height whilst manipulating vergence angle caused gross disruption to prehension, with DF mis-programming the reach component in a predictable manner. Notably, the evidence for DF's reliance on vergence distance information was obtained in a task where the targets were viewed at eye height. These experiments indicate that DF uses vertical gaze angle to gauge target distance in normal prehension and suggest that this extra-retinal cue may be a useful source of distance information for the human nervous system, especially where pictorial cues are impoverished.

Prehension and perception of size in left visual neglect.

Right hemisphere damaged patients with and without left visual neglect, and age-matched controls had objects of various sizes presented within left or right body hemispace. Subjects were asked to estimate the objects' sizes or to reach out and grasp them, in order to assess visual size processing in perceptual-experiential and action-based contexts respectively. No impairments of size processing were detected in the prehension performance of the neglect patients but a generalised slowing of movement was observed, associated with an extended deceleration phase. Additionally both patient groups reached maximum grip aperture relatively later in the movement than did controls. For the estimation task it was predicted that the left visual neglect group would systematically underestimate the sizes of objects presented within left hemispace but no such abnormalities were observed. Possible reasons for this unexpected null finding are discussed.

A test between two hypotheses and a possible third way for the control of prehension.

We used an obstacle avoidance task to test two opposing accounts of how the nervous system controls prehension. The visuomotor account supposes that the system independently controls the grip formation and transport phase of prehensile movements. In contrast, the digit channel hypothesis suggests that the system controls the thumb and finger more or less independently. Our data strongly favoured the traditional visuomotor channel hypothesis and demonstrated that the time taken to grasp an object in the presence of obstacles was well predicted by a Fitts' law relationship. We suggest a "thirdway" hypothesis in order to retain the advantages of the digit channel hypothesis within the visuomotor framework. The third-way hypothesis suggests that the nervous system selects a single digit to transport to the object. We speculate that the actual digit selected might depend upon attention and the nature of the prehension task. This hypothesis is able to account for most of the empirical findings unearthed by researchers investigating the control of prehension.

Improving the clinical diagnosis of personal neglect: a reformulated comb and razor test.

Beschin and Robertson (1997) devised a simple clinical test of left personal neglect, which characterises personal grooming behaviour according to the proportion of the total activity that is directed to the left side of the body. Although this test proved highly reliable, and more sensitive than prior diagnostic techniques, its formulation may yet be improved. The present paper reports a reanalysis of Beschin and Robertson's (1997) data, using additional control subjects, and a formula which characterises personal neglect as a lateral bias of behaviour rather than as a lateralised deficit. It is shown that this formula greatly enhances the test's sensitivity to the behavioural abnormalities of brain damaged patients, and it is recommended that this modification be adopted for the future diagnosis of personal neglect.