When the whole is more than the sum of the parts: Evidence from visuospatial neglect

One possible pathological mechanism underlying the rightward bisection error of right‐brain‐damaged patients with left spatial neglect is a leftward relaxation of the spatial representational medium. This view was originally based on the finding that patients with left neglect, required to extend horizontal segments, in order to double their original length, may exhibit a relative left overextension of the drawn lines ( Bisiach et al., 1994 ). We investigated this putative distortion of representational space using a 16 cm ‘line segmentation’ task (Experiment 1). Were the representation of space relaxed contralesionally, a progressive increase from right to left of the size of the drawn segments would be expected. Right‐brain‐damaged patients with left unilateral neglect (N=12) performed the segmentation task with no left versus right differences, as right‐brain‐damaged patients without neglect (N=8), and neurologically unimpaired control subjects (N=10), did. Experiments 2 and 3 explored the effects of sample length (1, 2, 4, and 8 cm), by which the 16 cm lines had to be segmented. Neglect patients produced longer left‐sided segments only for the 8 cm sample (i.e. half of the length of the segment). This set of experiments suggests an impairment in the segmentation task only with the larger (8 cm) sample, when a more global level of processing may be involved. Experiment 4 assessed this hypothesis by a ‘part/whole’ bisection task, using 8 cm lines, presented either embedded in a longer 16 cm line or in isolation. Neglect patients made a larger rightward bisection error when the segment was not embedded. The suggestion is made that the lateral distortion of the representation of space in neglect patients (i.e. a leftward relaxation of the spatial medium) concerns tasks where a more ‘global’ representation of the visual stimulus has to be set up. The different demands of the segmentation and bisection tasks are discussed.     

Pseudoneglect for the bisection of mental number lines

Patients with unilateral neglect of the left side bisect physical lines to the right whereas individuals with an intact brain bisect lines slightly to the left (pseudoneglect). Similarly, for mental number lines, which are arranged in a left-to-right ascending sequence, neglect patients bisect to the right. This study determined whether individuals with an intact brain show pseudoneglect for mental number lines. In Experiment 1, participants were presented with visual number triplets (e.g., 16, 36, 55) and determined whether the numerical distance was greater on the left or right side of the inner number. Despite changing the spatial configuration of the stimuli, or their temporal order, the numerical length on the left was consistently overestimated. The fact that the bias was unaffected by physical stimulus changes demonstrates that the bias is based on a mental representation. The leftward bias was also observed for sets of negative numbers (Experiment 2)—demonstrating not only that the number line extends into negative space but also that the bias is not the result of an arithmetic distortion caused by logarithmic scaling. The leftward bias could be caused by a rounding-down effect. Using numbers that were prone to large or small rounding-down errors, Experiment 3 showed no effect of rounding down. The task demands were changed in Experiment 4 so that participants determined whether the inner number was the true arithmetic centre or not. Participants mistook inner numbers shifted to the left to be the true numerical centre—reflecting leftward overestimation. The task was applied to 3 patients with right parietal damage with severe, moderate, or no spatial neglect (Experiment 5). A rightward bias was observed, which depended on the severity of neglect symptoms. Together, the data demonstrate a reliable and robust leftward bias for mental number line bisection, which reverses in clinical neglect. The bias mirrors pseudoneglect for physical lines and most likely reflects an expansion of the space occupied by lower numbers on the left side of the line and a contraction of space for higher numbers located on the right.     

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.     

Goal neglect and Spearman's g: competing parts of a complex task

In goal neglect, a person ignores some task requirement though being able to describe it. Goal neglect is closely related to general intelligence or C. Spearman's (1904) g (J. Duncan, H. Emslie, P. Williams, R. Johnson, & C. Freer, 1996). The authors tested the role of task complexity in neglect and the hypothesis that different task components in some sense compete for attention. In contrast to many kinds of attentional limits, increasing the real-time demands of one task component does not promote neglect of another. Neither does neglect depend on preparation for different possible events in a block of trials. Instead, the key factor is complexity in the whole body of knowledge specified in task instructions. The authors suggest that as novel activity is constructed, relevant facts, rules, and requirements must be organized into a "task model." As this model increases in complexity, different task components compete for representation, and vulnerable components may be lost. Construction of effective task models is closely linked to g.     

Hemispatial neglect and serial order in verbal working memory

Working memory refers to our ability to actively maintain and process a limited amount of information during a brief period of time. Often, not only the information itself but also its serial order is crucial for good task performance. It was recently proposed that serial order is grounded in spatial cognition. Here, we compared performance of a group of right hemisphere-damaged patients with hemispatial neglect to healthy controls in verbal working memory tasks. Participants memorized sequences of consonants at span level and had to judge whether a target consonant belonged to the memorized sequence (item task) or whether a pair of consonants were presented in the same order as in the memorized sequence (order task). In line with this idea that serial order is grounded in spatial cognition, we found that neglect patients made significantly more errors in the order task than in the item task compared to healthy controls. Furthermore, this deficit seemed functionally related to neglect severity and was more frequently observed following right posterior brain damage. Interestingly, this specific impairment for serial order in verbal working memory was not lateralized. We advance the hypotheses of a potential contribution to the deficit of serial order in neglect patients of either or both (1) reduced spatial working memory capacity that enables to keep track of the spatial codes that provide memorized items with a positional context, (2) a spatial compression of these codes in the intact representational space.     

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.     

Memory impairment following ruptured aneurysm of the anterior communicating artery

This study reports the case of a 42-year-old man who suffered a ruptured aneurysm of the anterior communicating artery. His memory capabilities were assessed after a considerable recovery period during which many of his memory deficits ameliorated. His scan revealed a left frontal lesion and many of his deficits were characteristic of frontal impairment. He was impaired on temporal discrimination, and he showed marked source forgetting. He also performed badly on the Brown-Peterson task, and we suggest that this is another task that may be characteristic of frontal impairment. In contrast, the patient showed normal or near normal performance on some memory tasks but not on others. It is concluded that the patient's frontal signs are similar to those found in Korsakoff's Syndrome, but that his memory impairment is qualitatively different from that encountered in patients with the amnesic syndrome.     

Misprojection of landmarks onto the spatial map

It has been suggested that neglect patients misrepresent the metric spatial relations along the horizontal axis (anisometry). The "fabric" of their internal spatial medium would be distorted in such a way that physically equal distances appear relatively shorter on the contralesional side (canonical anisometry). The case of GL, a 76-year-old lady with left neglect on visual search tasks, is presented. GL showed severe relative overestimation on the left (contralesional) side on two independent tasks evaluating the metrics of her internal representation. A qualitatively similar pattern was found in two out of 10 other neglect patients who performed the second task. This behavior cannot be accounted for by the canonical anisometry hypothesis. Nevertheless, GL produced a relative left overextension (underestimation) when trying to set the endpoints of a virtual line given its midpoint (Endpoints Task). An interpretation of these results is offered in terms of a misprojection of relevant landmarks onto the internal representation without assuming distortion of its "fabric."     

Does action make the link between number and space representation? Visuo-manual adaptation improves number bisection in unilateral neglect

If the visual world is artificially shifted by only 10 degrees, people initially experience difficulty in directing their actions toward visual goals, but then rapidly compensate the visual distortion. The consequence of such adaptation can be measured as visual and proprioceptive aftereffects, as well as by performance on pointing tasks without visual feedback. Recent work has shown that more cognitive deficits can be improved following prism adaptation in patients with unilateral neglect. Here we show that a short visuo-manual adaptation to prisms improves performance on a mental number-bisection task recently shown to be impaired in unilateral neglect. The association previously found between space and number representation (the mental number line) may thus be grounded in common action principles. Our results suggest that visuo-motor plasticity functionally links parietal areas involved in space and number representation.     

Pseudoneglect for the bisection of mental number lines

Patients with unilateral neglect of the left side bisect physical lines to the right whereas individuals with an intact brain bisect lines slightly to the left (pseudoneglect). Similarly, for mental number lines, which are arranged in a left-to-right ascending sequence, neglect patients bisect to the right. This study determined whether individuals with an intact brain show pseudoneglect for mental number lines. In Experiment 1, participants were presented with visual number triplets (e.g., 16, 36, 55) and determined whether the numerical distance was greater on the left or right side of the inner number. Despite changing the spatial configuration of the stimuli, or their temporal order, the numerical length on the left was consistently overestimated. The fact that the bias was unaffected by physical stimulus changes demonstrates that the bias is based on a mental representation. The leftward bias was also observed for sets of negative numbers (Experiment 2)--demonstrating not only that the number line extends into negative space but also that the bias is not the result of an arithmetic distortion caused by logarithmic scaling. The leftward bias could be caused by a rounding-down effect. Using numbers that were prone to large or small rounding-down errors, Experiment 3 showed no effect of rounding down. The task demands were changed in Experiment 4 so that participants determined whether the inner number was the true arithmetic centre or not. Participants mistook inner numbers shifted to the left to be the true numerical centre--reflecting leftward overestimation. The task was applied to 3 patients with right parietal damage with severe, moderate, or no spatial neglect (Experiment 5). A rightward bias was observed, which depended on the severity of neglect symptoms. Together, the data demonstrate a reliable and robust leftward bias for mental number line bisection, which reverses in clinical neglect. The bias mirrors pseudoneglect for physical lines and most likely reflects an expansion of the space occupied by lower numbers on the left side of the line and a contraction of space for higher numbers located on the right.     

A further test of sequential-sampling models that account for payoff effects on response bias in perceptual decision tasks

Recently, Diederich and Busemeyer (2006) evaluated three hypotheses formulated as particular versions of a sequential-sampling model to account for the effects of payoffs in a perceptual decision task with time constraints. The bound-change hypothesis states that payoffs affect the distance of the starting position of the decision process to each decision bound. The drift-rate-change hypothesis states that payoffs affect the drift rate of the decision process. The two-stage-processing hypothesis assumes two processes, one for processing payoffs and another for processing stimulus information, and that on a given trial, attention switches from one process to the other. The latter hypothesis gave the best account of their data. The present study investigated two questions: (1) Does the experimental setting influence decisions, and consequently affect the fits of the hypotheses? A task was conducted in two experimental settings--either the time limit or the payoff matrix was held constant within a given block of trials, using three different payoff matrices and four different time limits--in order to answer this question. (2) Could it be that participants neglect payoffs on some trials and stimulus information on others? To investigate this idea, a further hypothesis was considered, the mixture-of-processes hypothesis. Like the two-stage-processing hypothesis, it postulates two processes, one for payoffs and another for stimulus information. However, it differs from the previous hypothesis in assuming that on a given trial exactly one of the processes operates, never both. The present design had no effect on choice probability but may have affected choice response times (RTs). Overall, the two-stage-processing hypothesis gave the best account, with respect both to choice probabilities and to observed mean RTs and mean RT patterns within a choice pair.     

Spatial probability cuing and right hemisphere damage

In this experiment we studied statistical learning, inter-trial priming, and visual attention. We assessed healthy controls and right brain damaged (RBD) patients with and without neglect, on a simple visual discrimination task designed to measure priming effects and probability learning. All participants showed a preserved priming effect for item color. Contrary to healthy controls and RBD participants without neglect, RBD participants with neglect did not show positional priming and both RBD groups learned the underlying spatial probability distribution of target locations to a lesser degree. To see if the latter deficiency could be improved, we tested a patient with long standing chronic spatial neglect on three separate days and observed improved identification times for left sided, high probability, targets. In summary, we found preservation of priming per se in people with spatial neglect. However, this was only clearly demonstrable for color priming and not for positional priming. Associated with this impairment was a difficulty in learning the overall statistical structure of target locations. In a patient with severe persistent neglect we were able to demonstrate that the deficit in statistical learning was not absolute, as this subject improved his identification times for targets appearing in high probability regions of the test display.     

Visuospatial asymmetry in dual‐task performance after subacute stroke

Various authors have referred to an association between neglect and non‐spatial components of attention. It has been suggested that an increase in attentional load could exacerbate neglect symptoms and reveal subtle, well‐compensated neglect. In the present study, 21 RH and 22 LH subacute stroke patients and 20 controls performed a computerized single‐detection task (CVRT) and a dual task (CVRT‐D) combining the detection task with a driving simulation task. Omissions, reaction times (RTs) and RT asymmetries were analysed to investigate the influence of increasing attentional load on neglect symptoms. RT asymmetries were most pronounced in RH patients. Although a clear increase in RT asymmetries between CVRT and CVRT‐D was observed, the amount of increase did not differ between both patient groups. Within both patient groups, correlations between RT asymmetries and ipsilesional RTs as a measure of general attention were significant in the single task but not in the dual task, indicating that increased attentional load may result in different degrees of lateralized and general attentional problems. Half of the patients with neglect on the BIT (Behavioural Inattention Test) showed increased RT asymmetries from CVRT to CVRT‐D. Moreover, two LH and RH patients without neglect symptoms on the BIT and CVRT showed distinctively increased asymmetries in the CVRT‐D, fostering the idea of an emergence of subtle neglect under increased attentional load. Dual‐task performance may draw attention towards patients who, without obvious signs of neglect, may show visuospatial attention deficits in complex situations.     

Spatial probability cuing and right hemisphere damage

In this experiment we studied statistical learning, inter-trial priming, and visual attention. We assessed healthy controls and right brain damaged (RBD) patients with and without neglect, on a simple visual discrimination task designed to measure priming effects and probability learning. All participants showed a preserved priming effect for item color. Contrary to healthy controls and RBD participants without neglect, RBD participants with neglect did not show positional priming and both RBD groups learned the underlying spatial probability distribution of target locations to a lesser degree. To see if the latter deficiency could be improved, we tested a patient with long standing chronic spatial neglect on three separate days and observed improved identification times for left sided, high probability, targets. In summary, we found preservation of priming per se in people with spatial neglect. However, this was only clearly demonstrable for color priming and not for positional priming. Associated with this impairment was a difficulty in learning the overall statistical structure of target locations. In a patient with severe persistent neglect we were able to demonstrate that the deficit in statistical learning was not absolute, as this subject improved his identification times for targets appearing in high probability regions of the test display.     

Trunk orientation induces neglect-like lateral biases in covert attention

The purpose of this study was to resolve a paradox in the literature on the effects of body orientation on spatial attention. Neuropsychological studies have found that real or simulated trunk rotation relieves contralesional inattention in patients with unilateral neglect, suggesting that trunk orientation affects how attention is allocated to space. However, in two previous studies, trunk orientation did not affect spatial attention in other populations. In this study, we investigated the effects of trunk orientation on the performance of a covert attention task by neurologically intact adults. The covert attention task allowed the evaluation of the effects of trunk orientation on both the allocation of attention to space and the ability to shift that attention to new locations. As in previous research, trunk orientation did not affect participants' response times (RTs) to validly cued targets. However, rotating participants' trunks to the left increased their RTs to invalidly cued targets on the right and decreased their RTs to invalidly cued targets on the left. These results indicate that trunk orientation induces directional biases in the ability to shift attention. Thus, for intact participants, trunk rotation created lateral biases in the covert attention task similar to those seen in neglect patients.     

Position of the Egocentric Reference and Directional Arm Movements in Right-Brain-Damaged Patients

We asked 12 right-brain-damaged patients (6 with left neglect signs and 6 without left neglect signs) to perform a straight ahead pointing task and a visual detection task with lateralized motor response, in order to investigate the relationship between the position of the egocentric reference and response time and accuracy in producing lateralized arm movements. Results showed that there was no correlation between the position of the egocentric reference and neglect signs, nor between the position of the egocentric reference and the latencies to direct a motor response toward either side of space. These findings were interpreted within the context of egocentric hypotheses of neglect. In particular, it was suggested that attentional or intentional neglect signs cannot be considered as a direct consequence of an ipsilesional deviation of the egocentric reference.     

The influence of contralesional targets on the cancellation of ipsilesional targets in unilateral neglect

Neglect of contralesional stimuli and perseverative behavior on ipsilesional stimuli may co-occur on a target cancellation task. Our aim was to investigate whether the presence vs. absence of contralesional targets can modify perseveration on ipsilesional targets. We studied four right brain-damaged patients with left neglect and perseverative behavior on screening cancellation tasks. We compared their cancellation performance in two conditions: (i) targets equally distributed on both sides of space and (ii) targets confined to the right side. One patient showed no significant difference in perseveration between these two conditions; in contrast, three patients perseverated significantly more in condition (i) than in condition (ii). These results suggest that, at least in some patients, information from the 'neglected' targets is not completely lost, but rather it affects behavior in the ipsilesional side. The traditional dichotomy 'bad space-good space' would not apply here.     

Non-lateralised deficits of drawing production in hemispatial neglect

Spatially lateralised deficits that typically define the hemispatial neglect syndrome have been shown to co-occur with other non-lateralised deficits of attention, memory, and drawing. However even a simple graphic task involves multiple planning components, including the specification of drawing start position and drawing direction. In order to investigate the influence of these factors in neglect we presented patients with a circle-copying task, and specified the drawing start point. The ability to draw from the instructed location was strongly related to tests that measure constructional abilities, but not related to start point laterality. In contrast, the direction in which patients drew the circle was affected by start point laterality: patients with neglect were less likely to draw in a typical direction when the cue was on the affected side of space and this was strongly related to severity of the neglect. Patients with neglect consistently produced circles that were smaller than the model; however, the scaling was not affected by the laterality of the start point, nor was the proportion of drawings correctly started at the cue. These findings demonstrate the complex relationship between neglect and even the simplest test for the syndrome.     

Hemispheric asymmetries in cognitive modeling: connectionist modeling of unilateral visual neglect

Neglect is an acquired cognitive disorder characterized by a lack of processing of one side of a stimulus or representational space. There are hemispheric asymmetries in its cause and in its effects, but implemented computational models of neglect have tended not to incorporate this fact. The authors report a series of neural network simulations of the line-bisection task. They test the hypothesis that simple, neuroanatomically realistic principles of connectivity in the nervous system can produce emergent behaviors that capture a wide range of quantitative and qualitative data observed in neglect patients presenting with general visuospatial neglect. They demonstrate that exploring low-level architectural principles in implemented computational models is both a productive avenue of research and offers the most parsimonious explanations of behaviors observed in patients.     

Effects of attention and unilateral neglect on auditory stream segregation

Two pairs of experiments studied the effects of attention and of unilateral neglect on auditory streaming. The first pair showed that the build up of auditory streaming in normal participants is greatly reduced or absent when they attend to a competing task in the contralateral ear. It was concluded that the effective build up of streaming depends on attention. The second pair showed that patients with an attentional deficit toward the left side of space (unilateral neglect) show less stream segregation of tone sequences presented to their left than to their right ears. Streaming in their right ears was similar to that for stimuli presented to either ear of healthy and of brain-damaged controls, who showed no across-ear asymmetry. This result is consistent with an effect of attention on streaming, constrains the neural sites involved, and reveals a qualitative difference between the perception of left- and right-sided sounds by neglect patients.