Ipsilesional Stimuli Research Articles

Temporal feature integration in the right parietal cortex

When searching for a target presented among distractors by means of Rapid Serial Visual Presentation (RSVP), participants often report the stimulus that is preceding or following the target as being the target. These so-called temporal binding errors are accompanied by high levels of confidence so that participants are bemused with the mismatch between their perceptual experience and the actual presented stimulus. By contrast with spatial binding the neural basis for temporal binding errors remains unexplored. Previous neuropsychology studies using non-spatial selective attention tasks have shown that right temporo-parietal cortex is involved in the temporal deployment of attention. Here we investigated the neural basis of temporal binding in five patients with visual extinction whose lesions involved different cortical areas in the right hemisphere, including the temporo-parietal cortex. Patients made significantly more binding errors for contralesional than ipsilesional stimuli and more binding errors than healthy controls. Incorrect binding from distractors near to the target was the most common for both patients and controls. Eye movements did not contribute to the pattern of results. These results show that right hemisphere cortical areas contribute to the accurate temporal coding of visual features.

The role of the lateral PFC in inhibitory motor control

Event Abstract Back to Event The role of the lateral PFC in inhibitory motor control Ulrike Krämer1*, Anne-Kristin Solbakk2, Ingrid Funderud2, Marianne Lovstad2, Tor Endestad2 and Robert T. Knight1 1 University of California at Berkeley, United States 2 University Hospital Oslo, Norway Research on inhibitory motor control has implicated several prefrontal as well as subcortical and parietal regions in response inhibition. Yet, the exact functional role of prefrontal regions in inhibitory control, be it inhibition itself or rather attention or task-set representation, is still under debate. We investigated the influence of the lateral PFC on response inhibition by using cognitive electrophysiology in prefrontal lesion patients. Patients and age- and education-matched controls performed in a visual stop-signal task featuring lateralized stimuli, challenging either the intact or lesioned hemisphere. Participants also underwent a purely behavioral Go/Nogo task, which included a manipulation of inhibition difficulty (blocks with 50 vs. 80% go-trials). On the behavioral level, patients showed a tendency for a hemifield effect on the stop-signal reaction time (SSRT) with a slower SSRT for contra- compared to ipsilesional stimuli. Additionally, patients made more errors than controls in the Go/Nogo task, in particular in the more demanding condition. In controls, we observed after stop-stimuli an early lateralized negativity over parieto-occipital electrodes which was maximal contralateral to the visual stimulus. In the time-frequency domain, this was reflected in higher theta power and inter-trial coherence. Importantly, prefrontal patients showed a diminished negativity as well as theta power and coherence selectively over the lesioned hemisphere. Lesion patients also showed reduced frontal theta power compared to controls. The theta results can be explained with a PFC-dependent, attention-related modulation of visual processing which is diminished in patients and likely contributing to their impaired performance in the tasks. Funding: Supported by DFG Fellowship KR3691/1-1; NINDS Grants NS21135 and PO40813. Keywords: cognitive control, PFC Conference: XI International Conference on Cognitive Neuroscience (ICON XI), Palma, Mallorca, Spain, 25 Sep - 29 Sep, 2011. Presentation Type: Poster Presentation Topic: Poster Sessions: Performance Monitoring and Cognitive Control Citation: Krämer U, Solbakk A, Funderud I, Lovstad M, Endestad T and Knight RT (2011). The role of the lateral PFC in inhibitory motor control. Conference Abstract: XI International Conference on Cognitive Neuroscience (ICON XI). doi: 10.3389/conf.fnhum.2011.207.00249 Copyright: The abstracts in this collection have not been subject to any Frontiers peer review or checks, and are not endorsed by Frontiers. They are made available through the Frontiers publishing platform as a service to conference organizers and presenters. The copyright in the individual abstracts is owned by the author of each abstract or his/her employer unless otherwise stated. Each abstract, as well as the collection of abstracts, are published under a Creative Commons CC-BY 4.0 (attribution) licence (https://creativecommons.org/licenses/by/4.0/) and may thus be reproduced, translated, adapted and be the subject of derivative works provided the authors and Frontiers are attributed. For Frontiers’ terms and conditions please see https://www.frontiersin.org/legal/terms-and-conditions. Received: 21 Nov 2011; Published Online: 28 Nov 2011. * Correspondence: Dr. Ulrike Krämer, University of California at Berkeley, Berkeley, United States, umkraemer@gmail.com Login Required This action requires you to be registered with Frontiers and logged in. To register or login click here. Abstract Info Abstract The Authors in Frontiers Ulrike Krämer Anne-Kristin Solbakk Ingrid Funderud Marianne Lovstad Tor Endestad Robert T Knight Google Ulrike Krämer Anne-Kristin Solbakk Ingrid Funderud Marianne Lovstad Tor Endestad Robert T Knight Google Scholar Ulrike Krämer Anne-Kristin Solbakk Ingrid Funderud Marianne Lovstad Tor Endestad Robert T Knight PubMed Ulrike Krämer Anne-Kristin Solbakk Ingrid Funderud Marianne Lovstad Tor Endestad Robert T Knight Related Article in Frontiers Google Scholar PubMed Abstract Close Back to top Javascript is disabled. Please enable Javascript in your browser settings in order to see all the content on this page.

The grouping benefit in extinction: Overcoming the temporal order bias

Grouping between contra- and ipsilesional stimuli can alleviate the lateralised bias in spatial extinction ( Gilchrist, Humphreys, & Riddoch, 1996; Ward, Goodrich, & Driver, 1994). In the current study we demonstrate for the first time that perceptual grouping can also modulate the spatio/temporal biases in temporal order judgements affecting the temporal as well as the spatial coding of stimuli. Perceived temporal order was assessed by presenting two coloured letter stimuli in either hemi-field temporally segregated by a range of onset-intervals. Items were either identical (grouping condition) or differed in both shape and colour (non-grouping condition). Observers were required to indicate which item appeared second. Patients with visual extinction had a bias against the contralesional item appearing first, but this was modulated by perceptual grouping. When both items were identical in shape and colour the temporal bias against reporting the contralesional item was reduced. The results suggest that grouping can alter the coding of temporal relations between stimuli.

Asynchronous stimulus presentation in visual extinction: A psychophysical study

Patients showing visual extinction as a consequence of a unilateral brain lesion can correctly detect a single stimulus in either hemifield but fail to detect the contralesional stimulus (or at least process the stimulus less efficiently) when it is presented simultaneously with an ipsilesional stimulus. In an attempt to uncover the nature of the underlying deficit, some studies have manipulated the temporal characteristics of stimulus presentation. Contra- and ipsilesional stimuli with different stimulus onset asynchronies are typically used. In the present study, visual extinction was investigated in a group of left neglect patients (N=10) using a psychophysical paradigm with different stimulus onset asynchronies of target and distractor stimuli presented in different hemifields. Contrast thresholds for a target grating were determined with the target either in isolation or in the presence of an irrelevant distractor grating. When target and distractor gratings were presented simultaneously, neglect patients showed a significant extinction effect, i.e., a significant interference from the right hemifield distractor with left hemifield contrast sensitivity. When the right hemifield distractor preceded the left hemifield target stimulus by 250 ms, two different patterns of results were observed in the neglect patients. Five patients showed a significant improvement compared to the simultaneous presentation condition, five other patients showed a significant increase of the extinction effect. The results suggest that different underlying mechanisms, maybe due to different lesion locations, can cause extinction in neglect patients.

Horizontal and vertical dimensions of visual extinction: a theta burst stimulation study

After a lesion of the posterior parietal cortex (PPC), the perception of a contra-lesional stimulus in presence of a simultaneous, ipsilesional stimulus may be impaired, a phenomenon referred to as visual extinction. In the present study, visual extinction was transiently induced in healthy subjects by interfering with the function of the right PPC by means of continuous theta burst stimulation (TBS). We investigated to which extent the horizontal and vertical position of visual stimuli influenced the extinction rate. A single TBS train over the right PPC induced a significant increase of left visual extinctions of at least 30 min. Left visual extinction rate was higher when the left sided visual stimulus was presented at a more eccentric position on the horizontal axis (irrespective of right sided visual stimulus position) and in the lower part of the visual field. The results are discussed within the framework of current explanatory models and of putative inter- and intrahemispheric mechanisms directing visuospatial attention.

Preattentive surface and contour grouping in Kanizsa figures: Evidence from parietal extinction

Visual extinction commonly occurs after unilateral, parietal brain damage and manifests in a failure to identify contralesional stimuli when presented simultaneously with other, ipsilesional stimuli – but full awareness for single stimulus presentations. However, extinction can be substantially reduced when preattentive grouping operations link fragmentary items across hemifields into a coherent object. For instance, one study demonstrated preserved access to bilateral stimulus segments when these could be grouped to form a Kanizsa square [Mattingley, J. B., Davis, G., & Driver, J. (1997). Preattentive filling-in of visual surfaces in parietal extinction. Science, 275, 671–674]. Here, we investigated the relative contributions of distinct object attributes to the spared access in Kanizsa figure completion in extinction, by systematically varying the degree to which bilateral surface filling-in and contour interpolations group disparate items. We demonstrate that surface information can substantially reduce extinction, whereas contour completions showed comparably smaller influences. In summary, such graded influences of object attributes support recurrent models of grouping, first, linking fragmentary parts into coherent surfaces and, second, interpolating the precise boundaries.

Sustained interactions between perception and action in visual extinction and neglect: Evidence from sequential pointing

Interactions between perception and action were examined by assessing the effects of action programming on extinction and neglect. In an extension of prior work, effects of sequential motor programming were assessed under conditions in which attention was first directed to an ipsilesional stimulus. Despite pointing and reporting a stimulus on the ipsilesional side first, programming a second action to the contralesional side reduced the spatial deficit on report, improving the report of contralesional stimuli (relative to when the patient just pointed to the ipsilesional side) while decreasing the report of ipsilesional items. The data suggest that perception and action interact through motor feedback to early visual coding, helping a patient overcome a lack of visual awareness to contralesional stimuli. This is effective even when attention has to be disengaged from the ipsilesional side suggesting that motor programming decreases ipsilesional capture and exerts a sustained influence on perception.

Are drawing perseverations part of the neglect syndrome?

Unilateral neglect patients typically omit to cancel contralesional targets. Moreover, they can repeatedly cancel ipsilesional stimuli exhibiting what is termed ‘perseverative behavior’. Two alternative accounts of this behavior have been proposed. According to one of them, it is considered as integral to neglect and due either to a perceptual (allochiria), or a premotor (directional hypokinesia) pathological mechanism leading to the ipsilesional displacement of contralesional responses. According to the other one, perseverations are interpreted as the consequence of motor–control–disinhibition co-occurring with, although independent of, spatial neglect. We compared some crucial predictions of these two hypotheses on a group of 10 right-brain-damaged patients, eight with neglect and two without neglect, showing a perseverative behavior in both conventional and experimental cancellation tasks. In our experiment, the spatial location and the numerosity of targets were manipulated to obtain different degrees of horizontal alignment between targets on the left and on the right of the central vertical axis of the sheet. We found that ipsilesional perseverations were not influenced by left neglected targets and were not correlated to neglect severity. Additionally, perseverative errors were associated with right basal ganglia lesions rather than with presence of neglect. These findings support the view that two different pathological mechanisms might be involved in left spatial neglect and ipsilesional perseverative behavior.

Central perceptual load does not reduce ipsilesional flanker interference in parietal extinction.

In healthy individuals, filtering of distractors improves when the perceptual difficulty, or load, of a central task increases. Following an earlier study by Lavie and Robertson (2001), this study examined whether increasing the perceptual load of a visual discrimination task attenuates the influence of ipsilesional and contralesional distractors in patients with spatial extinction. The authors used a flanker task in which participants identified a central target letter flanked by congruent, incongruent, or neutral distractors in the left and right hemifields. Perceptual load was manipulated by positioning the central target letter above or below a hash mark (#) in the low-load condition, or a letter ("R") in the high-load condition. Target identification was significantly more difficult under high load than low load. Despite this difference in task difficulty, the interference from incongruent flankers was equivalent across the two load conditions, for both contralesional and ipsilesional flankers. Our results suggest that perceptual load does not attenuate the distracting influence of ipsilesional stimuli. Rather, selectivity is strongly influenced by the strength of representations in brain areas that code salience across the visual field.

A dissociation between visual and auditory hemi-inattention: Evidence from temporal order judgements

Patients with right hemisphere brain lesions often suffer from deficits in spatial attention that can be manifested in different sensory modalities. It has recently been claimed that a relationship (i.e., association) could exist between symptoms of hemi-inattention in different modalities, based on correlations between the results of visual and auditory clinical tests of neglect or extinction. However, it should be noted that the visual and auditory tasks varied greatly both in response type and level of sensitivity. Here, we have examined cross-modal associations in spatial attention deficits using a temporal order judgment task (TOJ) in which patients were required to identify which of two visual or auditory objects had appeared first. When compared to age and education matched control participants, the patients needed, on average, the contralesional stimulus to lead the ipsilesional stimulus to achieve the point of subjective simultaneity (PSS). No association between the degree of visual and auditory hemi-inattention was observed amongst the patients, suggesting that there is a certain degree of independence between the mechanisms subserving spatial attention across sensory modalities.

Competition between simultaneous stimuli modulated by location probability in hemispatial neglect

Many aspects of spatial neglect can be explained as arising from competition for attentional selection, with salient ipsilesional stimuli emerging as the winner more often than contralesional stimuli. The outcome of the competition, however, can be affected both by bottom-up perceptual factors such as the gestalt properties of the display and by top-down factors such as expectancy or stimulus blocking. This study examines whether the competition for attentional selection can be modulated by manipulating the probability of the target's location in hemispatial neglect. Five patients with left-sided hemispatial neglect and a group of control participants performed a visual target discrimination task. In equal probability blocks, the target appeared randomly in any of six possible horizontal locations (three left, three right) whereas in biased blocks, the target appeared in the mid-location on the left on 50% of the trials and in each of the other locations on 10% of the trials. The target appeared either alone or was accompanied by a distractor on the opposite side. The results showed that the spatial bias facilitated detection of all left-sided targets in the neglect group, but was more spatially specific in the control group. Furthermore, while distractors on either side interfered with target processing in both groups, the patterns differed across the visual field. Finally, the magnitude of facilitation due to the bias was greatest in the condition with the most inhibition, i.e. a left-sided target accompanied by a right-sided distractor in the neglect group. These data underscore the competitive push–pull relationship between different bottom-up and top-down attentional factors, particularly within neglect patients, in whom a strong ipsilesional attentional bias already exists.

Goal-driven selective attention in patients with right hemisphere lesions: how intact is the ipsilesional field?

Patients with right hemisphere (RH) lesions often display a spatial bias in attention towards the ipsilesional hemifield. The behavioural manifestations of this spatial bias are typically interpreted as reflecting increased or enhanced attention for stimuli within the 'intact' ipsilesional field, and impaired attentional functioning within the contralesional field. In the healthy brain, goal-driven and stimulus-driven attentional processes interact to determine which stimuli should be prioritized for selection. Although unilateral brain damage increases the relative attentional salience of stimuli within the ipsilesional field, it might also cause problems in filtering or attenuating task-irrelevant information. We examined whether goal-driven attention modulates the processing of ipsilesional and contralesional information in 6 patients with unilateral brain damage following RH stroke (5 male, 1 female; mean age 60.8 years) and a group of age and sex-matched controls. We used a flanker task in which participants made speeded judgements on a central target item (a coloured letter). On each trial the target was flanked by a coloured letter in the left and right hemifields. In separate blocks, participants were instructed to judge either the identity or the colour of the central target and to ignore the flankers. The flanker on one side could be congruent, incongruent or neutral with respect to the target, on either the letter or the colour dimension, whereas the flanker on the other side was always neutral on both dimensions. Healthy controls showed significant interference from incongruent flankers on either side. Crucially, however, this effect only occurred for the task-relevant dimension [F(2,10) = 24.60; P < 0.001]. For patients, however, both the task-relevant and task-irrelevant dimensions of ipsilesional flankers interfered with response times [task-relevant: F(2,10) = 7.50, P < 0.05; task-irrelevant: F(1,5) = 6.20, P < 0.05]. Conversely, contralesional flankers influenced response times only when the target and distractor were incongruent on the task-relevant dimension [F(2,10) = 4.85; P < 0.05]. Our findings demonstrate that following RH damage, goal-driven biases cannot constrain the processing of task-irrelevant features of ipsilesional stimuli. We speculate that a lateralized bias in spatial attention leads to unselective prioritization of all feature-based attributes of stimuli appearing within the ipsilesional hemifield, whether or not they are relevant to performance. Attentional selection for ipsilesional stimuli in disorders such as spatial neglect and extinction may not therefore be entirely normal, as previously assumed.

Reflexive Orienting in Spatial Neglect Is Biased towards Behaviourally Salient Stimuli

Patients with spatial neglect are impaired when detecting contralesional targets presented shortly after an ipsilesional cue. This "disengagement" deficit is believed to reflect reflexive orienting towards ipsilesional stimuli that is independent of behavioural goals. Here, we show that the extent of this spatial bias depends on the behavioural salience of ipsilesional stimuli. Healthy participants, brain-injured patients without neglect and neglect patients reacted to ipsilesional and contralesional visual targets. Prior to target presentation, a visual cue similar or dissimilar to the target was presented at target position or opposite the target. Although participants did not react to the similar cue, it had high behavioural salience since it shared features with the target stimulus. Neglect patients showed dramatically increased reaction times to contralesional targets, but only when these followed behaviourally relevant ipsilesional cues. No decrease of performance was observed with irrelevant cues. This performance pattern was not due to perceptual similarity, since the same effect was found when cue and target were semantically related but differed perceptually. Importantly, semantically related cues ceased to attract attention when they were defined as behaviourally irrelevant. These results show that neglect patients only orient attention reflexively towards ipsilesional stimuli with high behavioural salience.

Visuomotor links in awareness: evidence from extinction

In patients with extinction, ipsilesional stimuli may abolish awareness of contralesional stimuli. Explanations of extinction often assume a serial model of processing in which sensory competition and identification precedes the selection of responses. We tested the adequacy of this assumption by examining the effects of response variables on visual awareness in six patients using signal detection analysis. Ipsilesional stimuli modulated patients' response criteria in deciding whether a contralesional stimulus was a target, and response modality (verbal or motor) modulated patients' abilities to discriminate between contralesional targets and distractors. This pattern of input variables modulating response criteria and output variables modulating discriminability indicates the extent to which attentional and intentional systems are tightly intertwined, with bi-directional effects in producing visual awareness.

Role of disengagement failure and attentional gradient in unilateral spatial neglect – a longitudinal study

Purpose: To assess the importance of ‘disengagement failure’ and ‘attentional gradient’ in unilateral spatial neglect (USN) and in recovery from neglect.Method: Eight right-hemisphere-damaged stroke patients performed the standardized Behavioural-Inattention-Test battery for visual neglect, line-bisection tests, and two computerized reaction-time (RT) tasks: a variant of Posner's ‘Spatial-Cueing’ paradigm (with special emphasis on the magnitude of leftward disengagement time) and a signal-detection task (marking the spatial gradient of attention by the distribution of RTs to target stimuli in different spatial locations). The correlation between the different measures was assessed at two points in time, before and after a period of rehabilitation treatment.Results: A recovery pattern could be identified in both RT paradigms. However, the correlation between standard measures of neglect and performance on both, spatial-cueing and signal-detection tasks, was weak.Conclusion: Neither difficulty disengaging attention from an ipsilesional stimulus nor changes in the attentional gradient can fully explain the processes underlying USN and its recovery. A large interpersonal variance exists among USN patients in the expression of disengagement and other spatial-attention deficits. Hence, individual patients should be tested by measuring different factors known to play a role in USN. This information is crucial for assigning the appropriate treatment for each patient in accord with the specific deficit revealed.

Sensory and response contributions to visual awareness in extinction.

Brain-damaged patients may extinguish contralesional stimuli when ipsilesional stimuli are presented simultaneously. Most theories of extinction postulate that stimuli compete for pathologically limited attentional resources with a bias to process ipsilesional over contralesional stimuli. Implicit in this view is the idea that responses follow the outcome of an earlier competition between inputs. In the current study of two patients, we used signal detection analyses to test the hypothesis that response criteria and response modalities also contribute to visual awareness. We found that identification was more sensitive than detection in uncovering deficits of contralesional awareness. Extinction was worse with bilateral stimuli when the ipsilesional stimulus was identical or similar to the target than when it was dissimilar. This diminished awareness was more likely to reflect a shift towards more conservative responses rather than diminished discrimination of contralesional stimuli. By contrast, one patient was better able to discriminate contralesional stimuli when using his contralesional limb to indicate awareness of targets than when using his ipsilesional limb. These data indicate that the nature of stimuli can modulate response criteria and the motor response can affect the sensory discriminability. Sensory discrimination and response output are not organized in a simple serial manner. Rather, input and output parameters interact in complicated ways to produce visual awareness. Visual awareness itself appears to be the outcome of two bottlenecks in processing, one having to do with sensory processing that may be covert and the other having to do with decision making, which by definition is overt. Finally, we advocate the use of signal detection analyses in studies of extinction, a method that has been surprisingly neglected in this line of research.

Spatial-temporal anisometries following right parietal damage

Patients with right parietal damage often have a lateralized deficit of spatial attention. In addition to a spatial deficit, such patients have also been reported to have a non-spatial deficit in temporal processing. Here, we tested the hypothesis that these spatial and temporal deficits might be linked if the right temporal-parietal cortex is important in integrating spatial and temporal attention. In AF, a patient with an acute right temporal-parietal stroke, we replicated previous observations showing that he was biased to judge ipsilesional stimuli as occurring before contralesional stimuli. More importantly, for vertically aligned stimuli, AF more accurately judged the temporal order of successive ipsilesional than contralesional stimuli. Furthermore, his contralesional performance improved with stimuli with larger vertical separations. Taken together, these findings provide additional evidence that right temporal-parietal damage produces a processing refractory period for stimuli in contralesional space that extends in both space and time. These findings are in agreement with other studies that suggest that the right temporal-parietal cortex is important in integrating the where and when of stimuli.

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.

The cortical substrate of visual extinction.

Neuroimaging studies investigated the attentional systems of the human brain revealing two networks, one for voluntary allocation of attention and another for stimulus-driven attentional processes. Whereas lesions of the latter system were supposed to lead to spatial neglect, we show that such lesions rather are typical for the occurrence of visual extinction. Extinction describes the inability of brain-damaged patients to detect a contralesional target in the presence of a competing ipsilesional stimulus. In a sample of consecutively admitted patients with right hemisphere stroke, we found dissociable cortical substrates for spatial neglect and visual extinction. There was a surprising congruency between the typical lesion site in patients with extinction and the activation clusters found in previous neuroimaging studies of healthy subjects. The results show that the temporo-parietal junction (TPJ), considered to be a crucial part of the stimulus-driven attentional network, is the neural substrate of visual extinction.

The cortical substrate of visual extinction

Neuroimaging studies investigated the attentional systems of the human brain revealing two networks, one for voluntary allocation of attention and another for stimulus-driven attentional processes. Whereas lesions of the latter system were supposed to lead to spatial neglect, we show that such lesions rather are typical for the occurrence of visual extinction. Extinction describes the inability of brain-damaged patients to detect a contralesional target in the presence of a competing ipsilesional stimulus. In a sample of consecutively admitted patients with right hemisphere stroke, we found dissociable cortical substrates for spatial neglect and visual extinction. There was a surprising congruency between the typical lesion site in patients with extinction and the activation clusters found in previous neuroimaging studies of healthy subjects. The results show that the temporo-parietal junction (TPJ), considered to be a crucial part of the stimulus-driven attentional network, is the neural substrate of visual extinction.