Reaching toward a target viewed through laterally refracting prisms results in adaptation of both... more Reaching toward a target viewed through laterally refracting prisms results in adaptation of both visual and (limb) proprioceptive spatial representations. Common ways to measure adaptation after-effect are to ask a person to point straight ahead with their eyes closed ("manual straight ahead", MSA), or to a seen target using their unseen hand ("open-loop pointing", OLP). MSA measures changes in proprioception only, whereas OLP measures the combined visual and proprioceptive shift. The behavioural and neurological mechanisms of prism adaptation have come under scrutiny following reports of reduced hemispatial neglect in patients following this procedure. We present evidence suggesting that shifts in proprioceptive spatial representations induced by prism adaptation are larger following lesions to the intraparietal cortex-a brain region that integrates retinotopic visual signals with signals of eye position in the orbit and that is activated during prism adaptation. Six healthy participants and six patients with unilateral intraparietal cortex lesions underwent prism adaptation. After-effects were measured with OLP and MSA. Aftereffects of control participants were larger when measured with OLP than with MSA, consistent with previous research and with the additional contribution of visual shift to OLP after-effects. However, Patients' OLP shifts were not significantly different to their MSA shifts. We conclude that, for the patients, correction of pointing errors during prism adaptation involved proportionally more changes to arm proprioception than for controls. Since lesions to intraparietal cortex led to enhanced realignment of arm proprioceptive representations, our results indirectly suggest that the intraparietal cortex could be key for visual realignment.
Journal of Experimental Psychology: Human Perception and Performance, 1989
We measured manual reaction time in normal human subjects to confirm that an eccentric visual sig... more We measured manual reaction time in normal human subjects to confirm that an eccentric visual signal has a biphasic effect on covert attention and eye movements. First, it summons attention and biases a saccade toward the signal; a subsequent inhibition of return then slows responses to signals at that location. A temporal hemifield dominance for inhibition of return was shown; this finding coverges with observations in neurologic patients to suggest that it is mediated by midbrain pathways. Endogenous orienting of attention, from a central arrow cue, did not activate inhibition of return, whereas endogenous saccade preparation did so as effectively as an exogenous signal, even when no saccade was made. Inhibition of return is activated by midbrain oculomotor pathways and may function as a location "tagging" mechanism to optimize efficiency of visual search.
To investigate the role of the posterior thalamus in controlling voluntary and visually triggered... more To investigate the role of the posterior thalamus in controlling voluntary and visually triggered eye movements and ocular fixation. Methods: The latency to initiate saccades to peripheral targets (visually triggered) and in response to verbal commands (voluntary) was measured in three patients with unilateral lesions of the posterior thalamus, in normal controls, and in neurological controls with Parkinson's disease. On half the trials a fixation point offset simultaneously with target onset, and on half it remained visible. Results: Offset of the fixation point simultaneous with target onset decreased saccade latency for both voluntary and visually triggered eye movements in controls, but only for voluntary saccades in patients with thalamic lesions. Conclusions: These findings suggest that separate neural systems control fixation when making voluntary and visually triggered eye movements, and that the thalamus is involved in the control of fixation for visually triggered but not for voluntary saccades.
Reaching toward a target viewed through laterally refracting prisms results in adaptation of both... more Reaching toward a target viewed through laterally refracting prisms results in adaptation of both visual and (limb) proprioceptive spatial representations. Common ways to measure adaptation after-effect are to ask a person to point straight ahead with their eyes closed ("manual straight ahead", MSA), or to a seen target using their unseen hand ("open-loop pointing", OLP). MSA measures changes in proprioception only, whereas OLP measures the combined visual and proprioceptive shift. The behavioural and neurological mechanisms of prism adaptation have come under scrutiny following reports of reduced hemispatial neglect in patients following this procedure. We present evidence suggesting that shifts in proprioceptive spatial representations induced by prism adaptation are larger following lesions to the intraparietal cortex-a brain region that integrates retinotopic visual signals with signals of eye position in the orbit and that is activated during prism adaptation. Six healthy participants and six patients with unilateral intraparietal cortex lesions underwent prism adaptation. After-effects were measured with OLP and MSA. Aftereffects of control participants were larger when measured with OLP than with MSA, consistent with previous research and with the additional contribution of visual shift to OLP after-effects. However, Patients' OLP shifts were not significantly different to their MSA shifts. We conclude that, for the patients, correction of pointing errors during prism adaptation involved proportionally more changes to arm proprioception than for controls. Since lesions to intraparietal cortex led to enhanced realignment of arm proprioceptive representations, our results indirectly suggest that the intraparietal cortex could be key for visual realignment.
Space-and object-based attention components were examined in neurologically normal and parietal-l... more Space-and object-based attention components were examined in neurologically normal and parietal-lesion subjects, who detected a luminance change at 1 of 4 ends of 2 outline rectangles. One rectangle end was precued (75% valid); on invalid-cue trials, the target appeared at the other end of the cued rectangle or at 1 end of the uncued rectangle. For normals, the cost for invalid cues was greater for targets in the uncued rectangle, indicating an object-based component. Both rightand left-hemisphere patients showed costs that were greater for contralesional targets. For righthemisphere patients, the object cost was equivalent for contralesional and ipsilesional targets, indicating a spatial deficit, whereas the object cost for left-hemisphere patients was larger for contralesional targets, indicating an object deficit.
Using Eye Movements As an Experimental Probe of Brain Function - a Symposium in Honor of Jean Buttner-Ennever, 2008
This chapter was originally published in the book Progress in Brain Research, published by Elsevi... more This chapter was originally published in the book Progress in Brain Research, published by Elsevier, and the attached copy is provided by Elsevier for the author's benefit and for the benefit of the author's institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who know you, and providing a copy to your institution's administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution's website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier's
Proceedings of the National Academy of Sciences, 2001
To compare neural activity produced by visual events that escape or reach conscious awareness, we... more To compare neural activity produced by visual events that escape or reach conscious awareness, we used event-related MRI and evoked potentials in a patient who had neglect and extinction after focal right parietal damage, but intact visual fields. This neurological disorder entails a loss of awareness for stimuli in the field contralateral to a brain lesion when stimuli are simultaneously presented on the ipsilateral side, even though early visual areas may be intact, and single contralateral stimuli may still be perceived. Functional MRI and event-related potential study were performed during a task where faces or shapes appeared in the right, left, or both fields. Unilateral stimuli produced normal responses in V1 and extrastriate areas. In bilateral events, left faces that were not perceived still activated right V1 and inferior temporal cortex and evoked nonsignificantly reduced N1 potentials, with preserved face-specific negative potentials at 170 ms. When left faces were perce...
Proceedings of the National Academy of Sciences, 2009
We examined the contributions of the human pulvinar to goal directed selection of visual targets ... more We examined the contributions of the human pulvinar to goal directed selection of visual targets in 3 patients with chronic, unilateral lesions involving topographic maps in the ventral pulvinar. Observers completed 2 psychophysical tasks in which they discriminated the orientation of a lateralized target grating in the presence of vertically-aligned distracters. In experiment 1, where distracter contrast was varied while target contrast remained constant, the patients' contralesional contrast thresholds for discriminating the orientation of grating stimuli were elevated only when the task required selection of a visual target in the face of competition from a salient distracter. Attentional selectivity was restored in the patients in experiment 2 where target contrast was varied while distracter contrast remained constant. These observations provide the first evidence that the human pulvinar plays a necessary role in modulating physical saliency in attentional selection, and su...
Philosophical Transactions of the Royal Society of London. B, Biological Sciences, 1982
A peripheral visual cue in an empty field (1) often summons head or eyes, or both, (2) improves e... more A peripheral visual cue in an empty field (1) often summons head or eyes, or both, (2) improves efficiency at. the cued position while attention is directed to it, even without overt movements, and (3) reduces processing efficiency at the cued position once attention is withdrawn. We have studied the time course and the effects of mid-brain and cortical damage on these components of orienting. The facilitation arises from shifts in covert attention. In cases of mid-brain degeneration due to progressive supranuclear palsy, saccadic movements were abolished, while covert orienting still occurs. However, covert orienting was found to be delayed in directions in which eye movements were most affected, suggesting a role for mid-brain pathways in covert orienting. Parietal lesions can cause massive loss in detection contralateral to the lesion. This is especially true when attention has been directed to the opposite side. These findings relate aspects of covert orienting of attention to n...
The role of lateral prefrontal cortex in transducing perception into action was studied in 10 pat... more The role of lateral prefrontal cortex in transducing perception into action was studied in 10 patients with chronic, unilateral lesions. They identified colors in the center of a visual display, while a flanking, distractor color was presented simultaneously in either the ipsilesional or contralesional field. The flanker could be either the same color as the target, or incompatible with the correct response. The effects of compatible and incompatible flankers on reaction time (RT) served as a measure of response channel activation by the flanker. Flankers in the contralesional field influenced RT less than did those in the ipsilesional field. These results suggest that the lateral prefrontal cortex is involved in maintaining stimulus-response channels.
The current study investigated the role of the frontal eye fields (FEF) in the suppression of an ... more The current study investigated the role of the frontal eye fields (FEF) in the suppression of an unwanted eye movement ('oculomotor inhibition'). Oculomotor inhibition has generally been investigated using the antisaccade task, in which an eye movement to a task-relevant onset must be inhibited. Various lines of research have suggested that successful inhibition in the antisaccade task relies heavily on the FEF. Here, we tested whether the FEF are also involved in the oculomotor inhibition of reflexive saccades. To this end, we used the oculomotor capture task in which the to-be-inhibited element is task-irrelevant. Performance of four patients with lesions to the FEF was measured on both the antisaccade and oculomotor capture task. In both tasks, the majority of the patients made more erroneous eye movements to contralesional elements than to ipsilesional elements. One patient showed no deficits in the antisaccade task, which could be explained by the developmental origin of his lesion. While we confirm the role of the FEF in the inhibition of task-relevant elements, the current study also reveals that the FEF play a crucial role in the oculomotor inhibition of task-irrelevant elements.
Five patients with lesions involving intra-parietal cortex (IPCx) were tested in a rapid version ... more Five patients with lesions involving intra-parietal cortex (IPCx) were tested in a rapid version of the double step paradigm to investigate the role of the IPCx in the rapid, online, updating of a saccade program. Saccades were executed to a single target in either the contra-or the ipsilesional visual field. In two thirds of the trials, a step change in target position required that the saccade shifted to a new location within the same field but in the contra-or the ipsilesional direction, allowing us to investigate whether patients are able to update their saccade program given new exogenous information about the required endpoint of the saccade. This setup resulted in three types of initial saccades: saccades to the target on no-step trials, uncorrected saccades to the original target location on step trials and corrected saccades to the new target location on step trials. Furthermore, if the updating of the original eye movement program failed, patients performed a second saccade to the new target location that required a rapid error correction. The analysis of the double-step task on a group level indicated that latencies for all trial types were longer when saccades were directed to the contralesional versus the ipsilesional field. Furthermore, longer latencies were required for patients to initiate a corrective second saccade after making an uncorrected first saccade in their contralesional compared to ipsilesional field. There were no differences in the ultimate landing positions of the eye movements for such corrected saccades. These results reveal that deficits in updating of saccade programs only seem to be present if the updating must occur after the gaze has shifted to a new location, pointing to a role of intra-parietal cortex in the processes involved in updating information when the current reference frame has to be updated. In conclusion, the paradigm deployed in the current study allows for a refinement of the role of the intraparietal cortex in the updating of saccade programs.
The pulvinar nucleus of the thalamus, with its connections to visual areas and to frontal and par... more The pulvinar nucleus of the thalamus, with its connections to visual areas and to frontal and parietal oculomotor cortex, might serve as a nexus for integrating cortical control of voluntary eye movements with reflexive eye movements generated by the superior colliculus. To investigate this hypothesis, we tested five patients with a unilateral lesion of the pulvinar on the oculomotor capture paradigm. In this task, participants have to ignore a distractor item and make a saccade to a target in a visual search display. Results showed that the interference of the distractor was stronger when it was presented contralateral to their lesion compared to when it was presented in the ipsilesional visual field. These findings were confirmed by an additional single case experiment in which we measured saccade trajectory deviations as evoked by a single distractor. These results show that the pulvinar is involved in the successful influence of higher order signals (like our goals and intentions) on the guidance of our eye movements.
Monocular viewing conditions show an asymmetry between stimuli presented in the temporal and nasa... more Monocular viewing conditions show an asymmetry between stimuli presented in the temporal and nasal visual fields in their efficiency for automatically triggering eye saccades and grasping attention. For instance, observers free to make a saccade to one of two stimuli presented together orient preferentially to the temporal stimulus. Such naso-temporal asymmetry (NTA) has been assumed to reflect the asymmetry in the retinotectal pathway to the superior colliculus. We tested this hypothesis using S cone stimuli, which are invisible to the magnocellular and retinotectal pathways. The observed NTA in choice saccades to bilateral stimuli was no less present for S cone stimuli than for luminance stimuli. Additionally, the amplitude of the NTA can be enhanced when S cone signals are added to luminance signals. These results suggest that behavioral NTA in humans is not diagnostic of retinotectal mediation. Furthermore, we found no asymmetries in latency, suggesting that the NTA in saccade c...
Journal of Neurology, Neurosurgery & Psychiatry, 2004
To investigate the role of the posterior thalamus in controlling voluntary and visually triggered... more To investigate the role of the posterior thalamus in controlling voluntary and visually triggered eye movements and ocular fixation. Methods: The latency to initiate saccades to peripheral targets (visually triggered) and in response to verbal commands (voluntary) was measured in three patients with unilateral lesions of the posterior thalamus, in normal controls, and in neurological controls with Parkinson's disease. On half the trials a fixation point offset simultaneously with target onset, and on half it remained visible. Results: Offset of the fixation point simultaneous with target onset decreased saccade latency for both voluntary and visually triggered eye movements in controls, but only for voluntary saccades in patients with thalamic lesions. Conclusions: These findings suggest that separate neural systems control fixation when making voluntary and visually triggered eye movements, and that the thalamus is involved in the control of fixation for visually triggered but not for voluntary saccades.
Journal of Experimental Psychology: Human Perception and Performance, 1997
Object-based attention was examined in 2 split-brain patients. A precued object could move within... more Object-based attention was examined in 2 split-brain patients. A precued object could move within a visual field or cross the midiine to the opposite field. Normal individuals show an inhibition in detecting signals in the cued object whether it moves within or between fields. Both patients showed this effect when the cued object moved within a visual field. When it crossed the midiine into the opposite visual field, however, detection was faster in the cued box. These results reveal both facilitatory and inhibitory effects on attention that are object based and may last for several hundred milliseconds. However, the inhibition requires an intact corpus callosum for interhemispheric transfer, whereas the facilitation is transferred subeortically.
Maintaining a coherent percept of the visual scene while eye position continuously changes requir... more Maintaining a coherent percept of the visual scene while eye position continuously changes requires that saccades be accompanied by remapping of the visual environment. We studied saccadic remapping in patients with unilateral lesions in the intraparietal sulcus and healthy controls, using inhibition of return (IOR)—an inhibitory tag that enables efficient visual search. In healthy controls, IOR was found at both retinal and environmental locations of the cue, indicating that the inhibitory tag had been remapped into environmental coordinates. In contrast, right parietal patients demonstrated IOR only at the retinal location of the cue, indicating that the intraparietal sulcus is involved in remapping of the environment after eye movements to afford a stable, environmentally based reference frame. Note that patients did not show environmental IOR in either visual field. These results also suggest that this region may be the neural substrate for encoding inhibitory spatial tags in an...
Five patients with visual extinction following unilateral brain injury were briefly presented wit... more Five patients with visual extinction following unilateral brain injury were briefly presented with colored letters in either or both visual fields, and required to report and locate the colors or the shapes. On double simultaneous stimulation, they tended to miss the event contralateral to their lesion. This extinction was increased when the two stimuli were the same on the reported dimension, Similarity on the irrelevant dimension had no effect. These data suggest that extinguished colors and shapes may be correctly extracted by the visual system (when task-relaant) even though they are unavailable for verbal report. An analogy is made with the phenomena of “repetition blindness” in normal observers, and it is proposed that extinction may reflect failure in a token-individuation process for correctly extracted visual types.
It has been proposed that the mental representation of a graspable object involves not only a des... more It has been proposed that the mental representation of a graspable object involves not only a description of its visual properties but also encodings of the motor programs to act upon it [1]. Thus, observing a handle automatically primes the motor programs responsible for reaching and grasping it. Here, we provide neurological evidence that such action-related object features can bias visual selection. Two patients with visual extinction after right-parietal injury detected cups with left-or right-oriented handles, briefly displayed in either or both visual fields. People with this disorder have deficient awareness for stimuli toward the contralesional, left side of space, especially when competing stimuli appear further to the right [2]. This contralesional extinction was significantly reduced when cups had handles affording a left-hand grasp, even though no hand response was required. No effect was found when handles were replaced with patches equated for position, size, and mean luminance. These data suggest that action-related information may be correctly extracted by the visual system, even though they are unavailable for conscious report. It is proposed that an object affordance for grasping modulates attentional selection by activating specific motor schema that, in turn, enhance the competitive strength of that object representation. Results and Discussion Adaptive interactions with the environment require an intimate linkage between vision and action systems. Psychologist James J. Gibson [3] first introduced the notion of affordance, which refers to the property of an object or feature of the immediate environment that directly links perception and motor performance. There is now accumulating evidence, ranging from behavioral [1, 4] to physiological studies [5, 6], that simply viewing an object can partially activate possible actions toward it, even in the absence of explicit intentions to act. Thus, observing a handle automatically primes the motor programs for its reaching and grasping within a perceiver.
Cognitive, Affective, & Behavioral Neuroscience, 2004
In 3 patients with unilateral pulvinar lesions, we tested the pulvinar's role in selective attent... more In 3 patients with unilateral pulvinar lesions, we tested the pulvinar's role in selective attention processing. Each patient completed four variants of a flanker interference task in which they reported the color of a square of a specified size while ignoring an irrelevant flanker that appeared either contralesionally or ipsilesionally to the target. The main finding was that when target location was not known and target and flanker were associated with competing responses, reaction times to contralesional targets were longer than those to ipsilesional targets. Our findings suggest that pulvinar damage produces a contralesional deficit in response competition.
Reaching toward a target viewed through laterally refracting prisms results in adaptation of both... more Reaching toward a target viewed through laterally refracting prisms results in adaptation of both visual and (limb) proprioceptive spatial representations. Common ways to measure adaptation after-effect are to ask a person to point straight ahead with their eyes closed ("manual straight ahead", MSA), or to a seen target using their unseen hand ("open-loop pointing", OLP). MSA measures changes in proprioception only, whereas OLP measures the combined visual and proprioceptive shift. The behavioural and neurological mechanisms of prism adaptation have come under scrutiny following reports of reduced hemispatial neglect in patients following this procedure. We present evidence suggesting that shifts in proprioceptive spatial representations induced by prism adaptation are larger following lesions to the intraparietal cortex-a brain region that integrates retinotopic visual signals with signals of eye position in the orbit and that is activated during prism adaptation. Six healthy participants and six patients with unilateral intraparietal cortex lesions underwent prism adaptation. After-effects were measured with OLP and MSA. Aftereffects of control participants were larger when measured with OLP than with MSA, consistent with previous research and with the additional contribution of visual shift to OLP after-effects. However, Patients' OLP shifts were not significantly different to their MSA shifts. We conclude that, for the patients, correction of pointing errors during prism adaptation involved proportionally more changes to arm proprioception than for controls. Since lesions to intraparietal cortex led to enhanced realignment of arm proprioceptive representations, our results indirectly suggest that the intraparietal cortex could be key for visual realignment.
Journal of Experimental Psychology: Human Perception and Performance, 1989
We measured manual reaction time in normal human subjects to confirm that an eccentric visual sig... more We measured manual reaction time in normal human subjects to confirm that an eccentric visual signal has a biphasic effect on covert attention and eye movements. First, it summons attention and biases a saccade toward the signal; a subsequent inhibition of return then slows responses to signals at that location. A temporal hemifield dominance for inhibition of return was shown; this finding coverges with observations in neurologic patients to suggest that it is mediated by midbrain pathways. Endogenous orienting of attention, from a central arrow cue, did not activate inhibition of return, whereas endogenous saccade preparation did so as effectively as an exogenous signal, even when no saccade was made. Inhibition of return is activated by midbrain oculomotor pathways and may function as a location "tagging" mechanism to optimize efficiency of visual search.
To investigate the role of the posterior thalamus in controlling voluntary and visually triggered... more To investigate the role of the posterior thalamus in controlling voluntary and visually triggered eye movements and ocular fixation. Methods: The latency to initiate saccades to peripheral targets (visually triggered) and in response to verbal commands (voluntary) was measured in three patients with unilateral lesions of the posterior thalamus, in normal controls, and in neurological controls with Parkinson's disease. On half the trials a fixation point offset simultaneously with target onset, and on half it remained visible. Results: Offset of the fixation point simultaneous with target onset decreased saccade latency for both voluntary and visually triggered eye movements in controls, but only for voluntary saccades in patients with thalamic lesions. Conclusions: These findings suggest that separate neural systems control fixation when making voluntary and visually triggered eye movements, and that the thalamus is involved in the control of fixation for visually triggered but not for voluntary saccades.
Reaching toward a target viewed through laterally refracting prisms results in adaptation of both... more Reaching toward a target viewed through laterally refracting prisms results in adaptation of both visual and (limb) proprioceptive spatial representations. Common ways to measure adaptation after-effect are to ask a person to point straight ahead with their eyes closed ("manual straight ahead", MSA), or to a seen target using their unseen hand ("open-loop pointing", OLP). MSA measures changes in proprioception only, whereas OLP measures the combined visual and proprioceptive shift. The behavioural and neurological mechanisms of prism adaptation have come under scrutiny following reports of reduced hemispatial neglect in patients following this procedure. We present evidence suggesting that shifts in proprioceptive spatial representations induced by prism adaptation are larger following lesions to the intraparietal cortex-a brain region that integrates retinotopic visual signals with signals of eye position in the orbit and that is activated during prism adaptation. Six healthy participants and six patients with unilateral intraparietal cortex lesions underwent prism adaptation. After-effects were measured with OLP and MSA. Aftereffects of control participants were larger when measured with OLP than with MSA, consistent with previous research and with the additional contribution of visual shift to OLP after-effects. However, Patients' OLP shifts were not significantly different to their MSA shifts. We conclude that, for the patients, correction of pointing errors during prism adaptation involved proportionally more changes to arm proprioception than for controls. Since lesions to intraparietal cortex led to enhanced realignment of arm proprioceptive representations, our results indirectly suggest that the intraparietal cortex could be key for visual realignment.
Space-and object-based attention components were examined in neurologically normal and parietal-l... more Space-and object-based attention components were examined in neurologically normal and parietal-lesion subjects, who detected a luminance change at 1 of 4 ends of 2 outline rectangles. One rectangle end was precued (75% valid); on invalid-cue trials, the target appeared at the other end of the cued rectangle or at 1 end of the uncued rectangle. For normals, the cost for invalid cues was greater for targets in the uncued rectangle, indicating an object-based component. Both rightand left-hemisphere patients showed costs that were greater for contralesional targets. For righthemisphere patients, the object cost was equivalent for contralesional and ipsilesional targets, indicating a spatial deficit, whereas the object cost for left-hemisphere patients was larger for contralesional targets, indicating an object deficit.
Using Eye Movements As an Experimental Probe of Brain Function - a Symposium in Honor of Jean Buttner-Ennever, 2008
This chapter was originally published in the book Progress in Brain Research, published by Elsevi... more This chapter was originally published in the book Progress in Brain Research, published by Elsevier, and the attached copy is provided by Elsevier for the author's benefit and for the benefit of the author's institution, for non-commercial research and educational use including without limitation use in instruction at your institution, sending it to specific colleagues who know you, and providing a copy to your institution's administrator. All other uses, reproduction and distribution, including without limitation commercial reprints, selling or licensing copies or access, or posting on open internet sites, your personal or institution's website or repository, are prohibited. For exceptions, permission may be sought for such use through Elsevier's
Proceedings of the National Academy of Sciences, 2001
To compare neural activity produced by visual events that escape or reach conscious awareness, we... more To compare neural activity produced by visual events that escape or reach conscious awareness, we used event-related MRI and evoked potentials in a patient who had neglect and extinction after focal right parietal damage, but intact visual fields. This neurological disorder entails a loss of awareness for stimuli in the field contralateral to a brain lesion when stimuli are simultaneously presented on the ipsilateral side, even though early visual areas may be intact, and single contralateral stimuli may still be perceived. Functional MRI and event-related potential study were performed during a task where faces or shapes appeared in the right, left, or both fields. Unilateral stimuli produced normal responses in V1 and extrastriate areas. In bilateral events, left faces that were not perceived still activated right V1 and inferior temporal cortex and evoked nonsignificantly reduced N1 potentials, with preserved face-specific negative potentials at 170 ms. When left faces were perce...
Proceedings of the National Academy of Sciences, 2009
We examined the contributions of the human pulvinar to goal directed selection of visual targets ... more We examined the contributions of the human pulvinar to goal directed selection of visual targets in 3 patients with chronic, unilateral lesions involving topographic maps in the ventral pulvinar. Observers completed 2 psychophysical tasks in which they discriminated the orientation of a lateralized target grating in the presence of vertically-aligned distracters. In experiment 1, where distracter contrast was varied while target contrast remained constant, the patients' contralesional contrast thresholds for discriminating the orientation of grating stimuli were elevated only when the task required selection of a visual target in the face of competition from a salient distracter. Attentional selectivity was restored in the patients in experiment 2 where target contrast was varied while distracter contrast remained constant. These observations provide the first evidence that the human pulvinar plays a necessary role in modulating physical saliency in attentional selection, and su...
Philosophical Transactions of the Royal Society of London. B, Biological Sciences, 1982
A peripheral visual cue in an empty field (1) often summons head or eyes, or both, (2) improves e... more A peripheral visual cue in an empty field (1) often summons head or eyes, or both, (2) improves efficiency at. the cued position while attention is directed to it, even without overt movements, and (3) reduces processing efficiency at the cued position once attention is withdrawn. We have studied the time course and the effects of mid-brain and cortical damage on these components of orienting. The facilitation arises from shifts in covert attention. In cases of mid-brain degeneration due to progressive supranuclear palsy, saccadic movements were abolished, while covert orienting still occurs. However, covert orienting was found to be delayed in directions in which eye movements were most affected, suggesting a role for mid-brain pathways in covert orienting. Parietal lesions can cause massive loss in detection contralateral to the lesion. This is especially true when attention has been directed to the opposite side. These findings relate aspects of covert orienting of attention to n...
The role of lateral prefrontal cortex in transducing perception into action was studied in 10 pat... more The role of lateral prefrontal cortex in transducing perception into action was studied in 10 patients with chronic, unilateral lesions. They identified colors in the center of a visual display, while a flanking, distractor color was presented simultaneously in either the ipsilesional or contralesional field. The flanker could be either the same color as the target, or incompatible with the correct response. The effects of compatible and incompatible flankers on reaction time (RT) served as a measure of response channel activation by the flanker. Flankers in the contralesional field influenced RT less than did those in the ipsilesional field. These results suggest that the lateral prefrontal cortex is involved in maintaining stimulus-response channels.
The current study investigated the role of the frontal eye fields (FEF) in the suppression of an ... more The current study investigated the role of the frontal eye fields (FEF) in the suppression of an unwanted eye movement ('oculomotor inhibition'). Oculomotor inhibition has generally been investigated using the antisaccade task, in which an eye movement to a task-relevant onset must be inhibited. Various lines of research have suggested that successful inhibition in the antisaccade task relies heavily on the FEF. Here, we tested whether the FEF are also involved in the oculomotor inhibition of reflexive saccades. To this end, we used the oculomotor capture task in which the to-be-inhibited element is task-irrelevant. Performance of four patients with lesions to the FEF was measured on both the antisaccade and oculomotor capture task. In both tasks, the majority of the patients made more erroneous eye movements to contralesional elements than to ipsilesional elements. One patient showed no deficits in the antisaccade task, which could be explained by the developmental origin of his lesion. While we confirm the role of the FEF in the inhibition of task-relevant elements, the current study also reveals that the FEF play a crucial role in the oculomotor inhibition of task-irrelevant elements.
Five patients with lesions involving intra-parietal cortex (IPCx) were tested in a rapid version ... more Five patients with lesions involving intra-parietal cortex (IPCx) were tested in a rapid version of the double step paradigm to investigate the role of the IPCx in the rapid, online, updating of a saccade program. Saccades were executed to a single target in either the contra-or the ipsilesional visual field. In two thirds of the trials, a step change in target position required that the saccade shifted to a new location within the same field but in the contra-or the ipsilesional direction, allowing us to investigate whether patients are able to update their saccade program given new exogenous information about the required endpoint of the saccade. This setup resulted in three types of initial saccades: saccades to the target on no-step trials, uncorrected saccades to the original target location on step trials and corrected saccades to the new target location on step trials. Furthermore, if the updating of the original eye movement program failed, patients performed a second saccade to the new target location that required a rapid error correction. The analysis of the double-step task on a group level indicated that latencies for all trial types were longer when saccades were directed to the contralesional versus the ipsilesional field. Furthermore, longer latencies were required for patients to initiate a corrective second saccade after making an uncorrected first saccade in their contralesional compared to ipsilesional field. There were no differences in the ultimate landing positions of the eye movements for such corrected saccades. These results reveal that deficits in updating of saccade programs only seem to be present if the updating must occur after the gaze has shifted to a new location, pointing to a role of intra-parietal cortex in the processes involved in updating information when the current reference frame has to be updated. In conclusion, the paradigm deployed in the current study allows for a refinement of the role of the intraparietal cortex in the updating of saccade programs.
The pulvinar nucleus of the thalamus, with its connections to visual areas and to frontal and par... more The pulvinar nucleus of the thalamus, with its connections to visual areas and to frontal and parietal oculomotor cortex, might serve as a nexus for integrating cortical control of voluntary eye movements with reflexive eye movements generated by the superior colliculus. To investigate this hypothesis, we tested five patients with a unilateral lesion of the pulvinar on the oculomotor capture paradigm. In this task, participants have to ignore a distractor item and make a saccade to a target in a visual search display. Results showed that the interference of the distractor was stronger when it was presented contralateral to their lesion compared to when it was presented in the ipsilesional visual field. These findings were confirmed by an additional single case experiment in which we measured saccade trajectory deviations as evoked by a single distractor. These results show that the pulvinar is involved in the successful influence of higher order signals (like our goals and intentions) on the guidance of our eye movements.
Monocular viewing conditions show an asymmetry between stimuli presented in the temporal and nasa... more Monocular viewing conditions show an asymmetry between stimuli presented in the temporal and nasal visual fields in their efficiency for automatically triggering eye saccades and grasping attention. For instance, observers free to make a saccade to one of two stimuli presented together orient preferentially to the temporal stimulus. Such naso-temporal asymmetry (NTA) has been assumed to reflect the asymmetry in the retinotectal pathway to the superior colliculus. We tested this hypothesis using S cone stimuli, which are invisible to the magnocellular and retinotectal pathways. The observed NTA in choice saccades to bilateral stimuli was no less present for S cone stimuli than for luminance stimuli. Additionally, the amplitude of the NTA can be enhanced when S cone signals are added to luminance signals. These results suggest that behavioral NTA in humans is not diagnostic of retinotectal mediation. Furthermore, we found no asymmetries in latency, suggesting that the NTA in saccade c...
Journal of Neurology, Neurosurgery & Psychiatry, 2004
To investigate the role of the posterior thalamus in controlling voluntary and visually triggered... more To investigate the role of the posterior thalamus in controlling voluntary and visually triggered eye movements and ocular fixation. Methods: The latency to initiate saccades to peripheral targets (visually triggered) and in response to verbal commands (voluntary) was measured in three patients with unilateral lesions of the posterior thalamus, in normal controls, and in neurological controls with Parkinson's disease. On half the trials a fixation point offset simultaneously with target onset, and on half it remained visible. Results: Offset of the fixation point simultaneous with target onset decreased saccade latency for both voluntary and visually triggered eye movements in controls, but only for voluntary saccades in patients with thalamic lesions. Conclusions: These findings suggest that separate neural systems control fixation when making voluntary and visually triggered eye movements, and that the thalamus is involved in the control of fixation for visually triggered but not for voluntary saccades.
Journal of Experimental Psychology: Human Perception and Performance, 1997
Object-based attention was examined in 2 split-brain patients. A precued object could move within... more Object-based attention was examined in 2 split-brain patients. A precued object could move within a visual field or cross the midiine to the opposite field. Normal individuals show an inhibition in detecting signals in the cued object whether it moves within or between fields. Both patients showed this effect when the cued object moved within a visual field. When it crossed the midiine into the opposite visual field, however, detection was faster in the cued box. These results reveal both facilitatory and inhibitory effects on attention that are object based and may last for several hundred milliseconds. However, the inhibition requires an intact corpus callosum for interhemispheric transfer, whereas the facilitation is transferred subeortically.
Maintaining a coherent percept of the visual scene while eye position continuously changes requir... more Maintaining a coherent percept of the visual scene while eye position continuously changes requires that saccades be accompanied by remapping of the visual environment. We studied saccadic remapping in patients with unilateral lesions in the intraparietal sulcus and healthy controls, using inhibition of return (IOR)—an inhibitory tag that enables efficient visual search. In healthy controls, IOR was found at both retinal and environmental locations of the cue, indicating that the inhibitory tag had been remapped into environmental coordinates. In contrast, right parietal patients demonstrated IOR only at the retinal location of the cue, indicating that the intraparietal sulcus is involved in remapping of the environment after eye movements to afford a stable, environmentally based reference frame. Note that patients did not show environmental IOR in either visual field. These results also suggest that this region may be the neural substrate for encoding inhibitory spatial tags in an...
Five patients with visual extinction following unilateral brain injury were briefly presented wit... more Five patients with visual extinction following unilateral brain injury were briefly presented with colored letters in either or both visual fields, and required to report and locate the colors or the shapes. On double simultaneous stimulation, they tended to miss the event contralateral to their lesion. This extinction was increased when the two stimuli were the same on the reported dimension, Similarity on the irrelevant dimension had no effect. These data suggest that extinguished colors and shapes may be correctly extracted by the visual system (when task-relaant) even though they are unavailable for verbal report. An analogy is made with the phenomena of “repetition blindness” in normal observers, and it is proposed that extinction may reflect failure in a token-individuation process for correctly extracted visual types.
It has been proposed that the mental representation of a graspable object involves not only a des... more It has been proposed that the mental representation of a graspable object involves not only a description of its visual properties but also encodings of the motor programs to act upon it [1]. Thus, observing a handle automatically primes the motor programs responsible for reaching and grasping it. Here, we provide neurological evidence that such action-related object features can bias visual selection. Two patients with visual extinction after right-parietal injury detected cups with left-or right-oriented handles, briefly displayed in either or both visual fields. People with this disorder have deficient awareness for stimuli toward the contralesional, left side of space, especially when competing stimuli appear further to the right [2]. This contralesional extinction was significantly reduced when cups had handles affording a left-hand grasp, even though no hand response was required. No effect was found when handles were replaced with patches equated for position, size, and mean luminance. These data suggest that action-related information may be correctly extracted by the visual system, even though they are unavailable for conscious report. It is proposed that an object affordance for grasping modulates attentional selection by activating specific motor schema that, in turn, enhance the competitive strength of that object representation. Results and Discussion Adaptive interactions with the environment require an intimate linkage between vision and action systems. Psychologist James J. Gibson [3] first introduced the notion of affordance, which refers to the property of an object or feature of the immediate environment that directly links perception and motor performance. There is now accumulating evidence, ranging from behavioral [1, 4] to physiological studies [5, 6], that simply viewing an object can partially activate possible actions toward it, even in the absence of explicit intentions to act. Thus, observing a handle automatically primes the motor programs for its reaching and grasping within a perceiver.
Cognitive, Affective, & Behavioral Neuroscience, 2004
In 3 patients with unilateral pulvinar lesions, we tested the pulvinar's role in selective attent... more In 3 patients with unilateral pulvinar lesions, we tested the pulvinar's role in selective attention processing. Each patient completed four variants of a flanker interference task in which they reported the color of a square of a specified size while ignoring an irrelevant flanker that appeared either contralesionally or ipsilesionally to the target. The main finding was that when target location was not known and target and flanker were associated with competing responses, reaction times to contralesional targets were longer than those to ipsilesional targets. Our findings suggest that pulvinar damage produces a contralesional deficit in response competition.
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Papers by Robert Rafal