What Can Spatial Deficits Teach Us About Feature Binding and Spatial Maps?

The “binding problem” is discussed with reference to feature integration and visual search. Neuropsychological and neurophysiological findings support the existence of multiple visual areas within the primate cortex that respond to primary features of objects and spatial locations. Evidence from studies with hemineglect and Balint's syndrome supports the role of spatial attention in feature integration and demonstrates the necessity for an intact, explicitspatial representation. Some implicit spatial maps remain intact but are not sufficient to support the perception of properly bound features. The evidence suggests strong interactions between parietal spatial representations and temporal feature representations in feature integration.     

Remembering the past and imagining the future: a neural model of spatial memory and imagery

The authors model the neural mechanisms underlying spatial cognition, integrating neuronal systems and behavioral data, and address the relationships between long-term memory, short-term memory, and imagery, and between egocentric and allocentric and visual and ideothetic representations. Long-term spatial memory is modeled as attractor dynamics within medial-temporal allocentric representations, and short-term memory is modeled as egocentric parietal representations driven by perception, retrieval, and imagery and modulated by directed attention. Both encoding and retrieval/imagery require translation between egocentric and allocentric representations, which are mediated by posterior parietal and retrosplenial areas and the use of head direction representations in Papez's circuit. Thus, the hippocampus effectively indexes information by real or imagined location, whereas Papez's circuit translates to imagery or from perception according to the direction of view. Modulation of this translation by motor efference allows spatial updating of representations, whereas prefrontal simulated motor efference allows mental exploration. The alternating temporal-parietal flows of information are organized by the theta rhythm. Simulations demonstrate the retrieval and updating of familiar spatial scenes, hemispatial neglect in memory, and the effects on hippocampal place cell firing of lesioned head direction representations and of conflicting visual and ideothetic inputs.     

Crossover by Line Length and Spatial Location

It is well known that line length has a systematic influence on line bisection error in neglect. Most patients with neglect misbisect long lines on the same side of true center as their brain lesion but then cross over on short lines, misbisecting them on the opposite side (i.e., crossover by line length). What is less recognized is that the spatial location of lines relative to the viewer can similarly induce a crossover effect when one considers line bisection error scores that have been averaged across individual line lengths. Patients with right hemisphere injury and neglect classically make averaged line bisection errors that fall right of true center on lines located either at midline or to the left of the viewer; however, we observed that the averaged line bisection error can fall left of true center when lines are located to the right of the viewer (i.e., crossover by spatial location). We hypothesized that crossover by both line length and spatial location stem from systematic errors in magnitude estimation, i.e., perceived line length. We tested predictions based on this hypothesis by examining how the crossover effect by line length is altered by the spatial location of lines along a horizontal axis relative to the viewer. Participants included patients with unilateral lesions of the right and left cerebral hemispheres and age-appropriate normal subjects. All groups demonstrated a crossover effect by line length at the midline location but the effect was altered by placing lines to the right and left of the viewer. In particular, patients with right hemisphere injury and neglect crossed-over across a broader range of line lengths when the lines were located to the right of the viewer rather than at either midline or left of the viewer. It is proposed that mental representations of stimulus magnitude are altered in neglect, in addition to mental representations of space, and that traditional accounts of neglect can be enhanced by including the psychophysical concept of magnitude estimation.     

视觉空间伪忽视与表征伪忽视神经机制的异同

伪忽视是指个体存在的轻微偏左的不对称空间注意,而视觉空间伪忽视和表征伪忽视是其两种主要表现形式。研究初期,研究者认为两种伪忽视基于相同的注意定向左偏机制,但近年来研究发现,它们存在神经机制的差异。本文主要从两种伪忽视神经机制的异同出发,梳理近期研究结果,以期增进对伪忽视的理解。未来研究可以从认知时间进程角度或设计更为完善的研究范式进一步探讨这两种伪忽视神经机制的异同。     

Parietal versus temporal lobe components in spatial cognition: Setting the mid‐point of a horizontal line

Recent anatomo‐clinical correlation studies have extended to the superior temporal gyrus, the right hemisphere lesion sites associated with the left unilateral spatial neglect, in addition to the traditional posterior‐inferior‐parietal localization of the responsible lesion (supramarginal gyrus, at the temporo‐parietal junction). The study aimed at teasing apart, by means of repetitive transcranial magnetic stimulation (rTMS), the contribution of the inferior parietal lobule (angular gyrus versus supramarginal gyrus) and of the superior temporal gyrus of the right hemisphere, in making judgments about the mid‐point of a horizontal line, a widely used task for detecting and investigating spatial neglect. rTMS trains at 25 Hz frequency were delivered over the inferior parietal lobule (angular gyrus and supramarginal gyrus), the superior temporal gyrus and the anterior parietal lobe of the right hemisphere, in 10 neurologically unimpaired participants, performing a line bisection judgment task. rTMS of the inferior parietal lobule at the level of the supramarginal gyrus brought about a rightward error in the bisection judgment, ipsilateral to the side of the rTMS, with stimulation over the other sites being ineffective. The neural correlates of computing the mid‐point of a horizontal segment include the right supramarginal gyrus in the inferior parietal lobule and do not extend to the angular gyrus and the superior temporal gyrus. These rTMS data in unimpaired subjects constrain the evidence from lesion studies in brain‐damaged patients, emphasizing the major role of a subset of relevant regions.     

Spatial hemineglect in humans

The term `spatial hemineglect' refers collectively to disorders of spatial cognition, which concern specific sectors of space with reference to a given coordinate system. Patients with cerebral lesions involving the posterior-inferior parietal and the premotor cortex, most often in the right hemisphere, sometimes fail to explore the extra-personal and personal sectors of space contralateral to the side of the lesion, are not aware of stimuli presented in these portions of space, or of contralateral body parts and their disordered function. In addition to these negative signs patients may also show positive pathological manifestations, such as avoidance or withdrawal from the contralateral side of space, and delusional views concerning contralateral body parts. The many varieties of this disorder can occur in dissociated forms, suggesting a multifaceted organization of the internal representation of space, of spatial attention, and of their neural correlates. Many manifestations of hemineglect are modulated in a similar fashion by specific sensory stimulation that also affects visuo-motor processes in normal subjects. This ongoing sensory modulation might update the internal representations of space in a continuously changing environment, and contribute to the formation, around the vertical orientation of gravity, of our subjective unitary experience of space.     

Exaggerated leftward bias in the mental number line of patients with schizophrenia

Several visuo-motor tasks can be used to demonstrate biases towards left hemispace in schizophrenic patients, suggesting a minor right hemineglect. Recent studies in neglect patients used a new number bisection task to highlight a lateralized defect in their visuo-spatial representation of numbers. To test a possible lateralized representational deficit in schizophrenia, we used the number bisection task in 11 schizophrenic patients compared to 11 healthy controls. Participants were required to orally indicate the central number of an interval orally presented. Whereas healthy subjects showed no significant bias, schizophrenic patients presented a significant leftward bias. Therefore, these results suggest an impairment in higher order representations of the number space in patients with schizophrenia, an impairment that is qualitatively similar to the deficit described in neglect patients.     

Systematic analysis of deficits in visual attention

A variety of impairments in visual attention can follow damage to the brain. The authors develop systematic methods for analyzing such impairments in terms of C. Bundesen's (1990) Theory of Visual Attention and apply these in a group of 9 patients with parietal lobe lesions and variable spatial neglect. In whole report, patients report letters from brief, vertical arrays in left or right visual field. The results show substantial, largely bilateral impairments in processing capacity, implying a major nonlateralized aspect to neglect. In partial report, arrays contain 1 or 2 letters in red and/or green. The task is to report only those letters in a specified target color. In addition to the expected bias against left-sided letters, patients show striking, bilateral preservation of top-down control, or attentional priority for targets. The results show how differentiation of attentional impairments can be informed by a theory of normal function.     

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

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

The interaction of spatial reference frames and hierarchical object representations: evidence from figure copying in hemispatial neglect

In copying or drawing a figure, patients with hemispatial neglect following right parietal lobe lesions typically produce an adequate representation of parts on the right of the figure while omitting the corresponding features on the left. The neglect of information occupying contralateral locations is influenced by multiple spatial reference frames and by the hierarchical structure of the object(s) in the figure. The present work presents a computational characterization of the interaction among these influences to account for the way in which neglect manifests in copying. Empirical data are initially collected from brain-damaged and normal control subjects during two figure-copying tasks in which the hierarchical complexity and orientation of the displays to be copied are manipulated. In the context of the model, neglect is simulated by a “lesion” (monotonic drop-off along gradient from right to left) that can affect performance in both object- and viewer-centered reference frames. The effect of neglect in both these frames, coupled with the hierarchical representation of the object(s), provides a coherent account of the copying behavior of the patients and may be extended to account for the copying performance of other patients across a range of objects and scenes.     

Cognitive rehabilitation: attention and neglect

Cognitive neuroscience can make a significant contribution towards the development of a scientific basis for the practice of brain rehabilitation. Though rehabilitation is a vast worldwide industry, there is little scientific basis for the training and therapy that are designed to help damaged brain circuits to recover. The systematic application of cognitive neuroscience models to rehabilitation can not only foster better, more theoretically grounded rehabilitation, but the models themselves can be tested and modified by data generated in rehabilitation-oriented research. The example of unilateral spatial neglect is used here to show how non-intuitive but clinically tractable methods can emerge out of systematic application of cognitive neuroscience to the problem of how to foster dynamic change and recovery in the damaged brain. Examples are given of recently developed rehabilitation methods for unilateral spatial neglect that are both derived from theoretical models of cognitive function, and that feed back into these models. These include dorsal–ventral stream interactions, perceptuo–motor interactions, interhemispheric inhibitory dynamics, and arousal–spatial attention interactions. It will be to the mutual benefit of basic cognitive neuroscience and rehabilitation if this type of research is expanded into other domains of cognitive function, in which similar theory–practice interactions exist.     

Visuospatial Neglect: The Ultimate Deconstruction?

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

Rightward collisions and their association with pseudoneglect

Whereas right parietal damage can result in left hemineglect, the general population shows a subtle neglect of the right hemispace-known as pseudoneglect. A recent study has demonstrated that people collide to the right more often and attributed this bias to pseudoneglect. [Nicholls, M. E. R., Loftus, A., Meyer, K., & Mattingley, J.B. (2007). Things that go bump in the right: The effect of unimanual activity on rightward collisions. Neuropsychologia, 45, 1122-1126]. Nicholls examined the effect of unimanual activation by requiring participants to fire projectiles at a target whilst walking and found that the rightward bias was exaggerated or reversed when the left and right hands were active, respectively. However, the act of aiming at a target may have inadvertently biased walking trajectory to the right. The current study addressed this issue by requiring participants (n=149) to walk through a narrow doorway three times whilst entering text into a phone using the (a) left, (b) right or (c) both hands. Despite the fact that entering text into a phone should produce no rightward bias, participants bumped to the right more often. Unlike previous research, no effect of unimanual activation was observed. This lack of effect was attributed to the smaller hand movements for entering numbers compared to firing a toy gun. Finally, this study showed an association for the first time between biases in observable bumping and line bisection performance-suggesting that unilateral bumping is related to pseudoneglect.     

3-D vision and figure-ground separation by visual cortex

A neural network theory of three-dimensional (3-D) vision, called FACADE theory, is described. The theory proposes a solution of the classical figure-ground problem for biological vision. It does so by suggesting how boundary representations and surface representations are formed within a boundary contour system (BCS) and a feature contour system (FCS). The BCS and FCS interact reciprocally to form 3-D boundary and surface representations that are mutually consistent. Their interactions generate 3-D percepts wherein occluding and occluded object parts are separated, completed, and grouped. The theory clarifies how preattentive processes of 3-D perception and figure-ground separation interact reciprocally with attentive processes of spatial localization, object recognition, and visual search. A new theory of stereopsis is proposed that predicts how cells sensitive to multiple spatial frequencies, disparities, and orientations are combined by context-sensitive filtering, competition, and cooperation to form coherent BCS boundary segmentations. Several factors contribute to figure-ground pop-out, including: boundary contrast between spatially contiguous boundaries, whether due to scenic differences in luminance, color, spatial frequency, or disparity-partially ordered interactions from larger spatial scales and disparities to smaller scales and disparities; and surface filling-in restricted to regions surrounded by a connected boundary. Phenomena such as 3-D pop-out from a 2-D picture, Da Vinci stereopsis, 3-D neon color spreading, completion of partially occluded objects, and figure-ground reversals are analyzed. The BCS and FCS subsystems model aspects of how the two parvocellular cortical processing streams that join the lateral geniculate nucleus to prestriate cortical area V4 interact to generate a multiplexed representation of Form-And-Color-And-DEpth, orfacade, within area V4. Area V4 is suggested to support figure-ground separation and to interact with cortical mechanisms of spatial attention, attentive object learning, and visual search. Adaptive resonance theory (ART) mechanisms model aspects of how prestriate visual cortex interacts reciprocally with a visual object recognition system in inferotemporal (IT) cortex for purposes of attentive object learning and categorization. Object attention mechanisms of the What cortical processing stream through IT cortex are distinguished from spatial attention mechanisms of the Where cortical processing stream through parietal cortex. Parvocellular BCS and FCS signals interact with the model What stream. Parvocellular FCS and magnocellular motion BCS signals interact with the model Where stream. Reciprocal interactions between these visual, What, and Where mechanisms are used to discuss data about visual search and saccadic eye movements, including fast search of conjunctive targets, search of 3-D surfaces, selective search of like-colored targets, attentive tracking of multielement groupings, and recursive search of simultaneously presented targets.     

Paradoxical facilitation of attention in healthy humans

Transcranial magnetic stimulation (TMS)-induced virtual lesions in healthy subjects can be used to test neurofunctional models of disease. The interhemispheric rivalry model of heminglect is well suited for such investigations, as simple predictions derived from clinical data can be tested without the caveats normally associated with lesion studies. One of these predictions is that release from contralateral inhibition should lead to increased parietal responsiveness, which, in turn, would enhance spatial attention. Here, we detail studies showing TMS-induced paradoxical functional facilitation of attention in healthy individuals and highlight their contribution to the understanding and treatment of neglect syndromes.     

Attention accesses multiple reference frames: evidence from visual neglect

Research with normal participants has demonstrated that mechanisms of selective attention can simultaneously gain access to internal representations of spatial information defined with respect to both location- and object-based frames of reference. The present study demonstrates that patients with unilateral spatial neglect following a right-hemisphere lesion are poorer at detecting information on the contralateral left side in both location- and object-based spatial coordinates simultaneously. Moreover, the extent of the neglect is modulated by the probability of a target's appearing in either reference frame; as the probability of sampling a target in a particular frame of reference increases, so does the severity of neglect in the frame. These findings suggest that attention can be flexibly and strategically assigned to a reference frame depending on the contingencies of the task.     

Spatial neglect, Balint-Homes' and Gerstmann's syndrome, and other spatial disorders

Brain-damaged patients with lesion or dysfunction involving the parietal cortex may show a variety of neuropsychological impairments involving spatial cognition. The more frequent and disabling deficit is the syndrome of unilateral spatial neglect that, in a nutshell, consists in a bias of spatial representation and attention ipsilateral to of extrapersonal, personal (ie, the body) space, or both, toward the side of the hemispheric lesion. The deficit is more frequent and severe after damage to the right hemisphere, involving particularly the posterior-inferior parietal cortex at the temporo-parietal junction. Damage to these posterior parietal regions may also impair visuospatial short-term memory, which may be associated with and worsen spatial neglect. The neural network supporting spatial representation, attention and short-term memory is, however, more extensive, including the right premotor cortex. Also disorders of drawing and building objects (traditionally termed constructional apraxia) are a frequent indicator of posterior parietal damage in the left and in the right hemispheres. Other less frequent deficits, which, however, have a relevant localizing value, include optic ataxia (namely, the defective reaching of visual objects, in the absence of elementary visuo-motor impairments), which is typically brought about by damage to the superior parietal lobule. Optic ataxia, together with deficits of visual attention, of estimating distances and depth, and with apraxia of gaze, constitutes the severely disabling Balint-Holmes' syndrome, which is typically associated with bilateral posterior parietal and occipital damage. Finally, lesions of the posterior parietal lobule (angular gyrus) in the left hemisphere may bring about a tetrad of symptoms (left-right disorientation, acalculia, finger agnosia, and agraphia) termed Gerstmann's syndrome, that also exists in a developmental form.     

Posterior parietal cortex as part of a neural network for directed attention in rats

A rodent model of directed attention has been developed based upon behavioral analysis of contralateral neglect, pharmacological manipulations, and anatomical analysis of neural circuitry. In each of these three domains the rodent model exhibits striking similarities to humans. We hypothesize that there is a specific thalamo-cortical-basal ganglia network that subserves spatial attentional functions. Key components of this network are medial agranular and posterior parietal cortex, dorsocentral striatum, and the lateral posterior thalamic nucleus. Several issues need to be addressed before we can hope to realistically understand or model the functions of this network. Among these are the roles of medial versus lateral posterior parietal cortex; cholinergic mechanisms in attention; interhemispheric interactions; the role of synchronous firing at the cortical, striatal, and thalamic levels; interactions between cortical and thalamic projections to the striatum; interactions between cortical and nigral inputs to the thalamus; the role of collicular inputs to the lateral posterior thalamic nucleus; the role of cerebral cortex versus superior colliculus in driving the motor output expressed as orienting behavior during directed attention; the extent to which the circuitry we describe for directed attention also plays a role in other forms of attention.     

Interactions of space and language: Insights from the neglect syndrome

The current article speculates about the interrelations between space and language from the perspective of the neglect syndrome. Various conflicting findings are reviewed and the effect of cultural factors is strongly emphasised. Current scientific literature describes relationships between space and language; however, the available data render it difficult to conclude whether it is the primacy of spatial representations or the complex two‐way interaction between linguistic and spatial representations that determines the nature of cognitive processing. It is hoped that studying the neglect syndrome will provide a solution to this problem. Finally, directions for future research are provided that requires bridging the gap between cognitive theorising and neuroscientific approaches.