The contribution of difficulty of an irrelevant task to task conflict.

In the standard colour-word Stroop task, participants are presented with colour words and required to respond to their colour while ignoring their meaning. Two types of conflict might occur in such experiments: information conflict and task conflict. Information conflict reflects the processing of two contradicting pieces of information and is indicated by shorter reaction times (RTs) in congruent than in incongruent trials. Task conflict reflects the additional effort associated with performing two tasks, as opposed to one, and is indicated by shorter RTs in neutral trials than in congruent trials (termed reverse facilitation). While information conflict is commonly seen in Stroop and Stroop-like tasks, task conflict is rarely observed. In the present study, participants were presented with coloured segments that, by applying Gestalt principles, could be perceived as colour words. We found that incongruent trials were slower than congruent trials, suggesting that participants successfully perceived the colour words, which led to involuntary reading. In addition, reversed facilitation was found so that neutral trials (i.e., trials that only consist of one task) were faster than congruent trials (as well as incongruent trials; both consist of two tasks). The presence of both interference from the incongruent trials and reverse facilitation suggests that involuntary reading could also occur in scenarios requiring cognitive effort.

Top-down effect on pupillary response: Evidence from shape from shading.

Shaded 2D images often create an illusion of depth, due to the shading information and assumptions regarding the location of the light source. Specifically, 2D images that are lighter on top usually appear convex while images that are darker on top, usually appear concave, reflecting the assumption that light is coming from above. The process of recovering the 3D shape of a shaded image is called Shape from Shading. Here we examined whether the pupil responds to the illusion of depth in a shape from shading task. In three experiments we show that pupil size is affected by the percept of depth, so that it dilates more when participants perceive the stimulus as concave, compared to when they perceive it as convex. This only happens if participants make a judgment regarding the shape of the stimulus or when they view it passively but are aware of the different shapes. No differences in pupil size were found with passive viewing if participants were not aware of the illusion, suggesting that some aspects of shape from shading require attention. All stimuli were equiluminant, and the percept of depth was created by manipulating the orientation of the shading, so that changes in pupil size could not be accounted by changes in the amount of light in the image. We posit, and confirmed it in a behavioral control experiment, that the perception of depth is translated to a subjective perception of darkness, due to the "darker is deeper" heuristic and conclude that the pupillary physiological response reflects the subjective perception of light.

3D shape-from-shading relies on a light source prior that does not change with age.

The light-from-above prior enables observers to infer an object's three-dimensional shape-from-shading information. Young, Western adults implicitly assume the light source is placed not only above, but also to the left of, the observer. Previous evidence reached conflicting conclusions regarding the development of the assumed light source direction. In the present study, we measured the light source prior cross-sectionally in children aged 5-11 years, using an explicit shape judgement task. The light-from-above prior, and the left bias, were present as soon as children became sensitive to shading information, regardless of their age. Global processing preference was not related to the ability to perform the task. Similarly, scanning habits, as measured by reading proficiency and starting position in a cancellation task, were not related to the magnitude of the left bias. Children's ability to report shape-from-shading judgements increased with age, but age did not affect the direction of light priors. Thus, we concluded that the development of the light-from-above prior and leftward bias do not require an extended maturation period, but rather the direction of the light-source priors may be developmentally stable once measurable.

Dimensionally Specific Capture of Attention: Implications for Saliency Computation.

Observers automatically orient to a sudden change in the environment. This is demonstrated experimentally using exogenous cues, which prioritize the analysis of subsequent targets appearing nearby. This effect has been attributed to the computation of saliency, obtained by combining features specific signals, which then feed back to drive attention to the salient location. An alternative possibility is that cueing directly effects target-evoked sensory responses in a feed-forward manner. We examined the effects of luminance and equiluminant color cues in a dual task paradigm, which required both a motion and a color discrimination. Equiluminant color cues improved color discrimination more than luminance cues, but luminance cues improved motion discrimination more than equiluminant color cues. This suggests that the effects of exogenous cues are dimensionally specific and may not depend entirely on the computation of a dimension general saliency signal.

Human Parahippocampal Cortex Supports Spatial Binding in Visual Working Memory.

Studies investigating the functional organization of the medial temporal lobe (MTL) suggest that parahippocampal cortex (PHC) generates representations of spatial and contextual information used by the hippocampus in the formation of episodic memories. However, evidence from animal studies also implicates PHC in spatial binding of visual information held in short term, working memory. Here we examined a 46-year-old man (P.J.), after he had recovered from bilateral medial occipitotemporal cortex strokes resulting in ischemic lesions of PHC and hippocampal atrophy, and a group of age-matched healthy controls. When recalling the color of 1 of 2 objects, P.J. misidentified the target when cued by its location, but not shape. When recalling the position of 1 of 3 objects, he frequently misidentified the target, which was cued by its color. Increasing the duration of the memory delay had no impact on the proportion of binding errors, but did significantly worsen recall precision in both P.J. and controls. We conclude that PHC may play a crucial role in spatial binding during encoding of visual information in working memory.

Aging changes 3D perception: Evidence for hemispheric rebalancing of lateralized processes.

When judging the 3D shape of a shaded image, young observers assume that the light source is placed above and to the left. This leftward bias has been attributed to hemispheric lateralization or experiential factors. Since aging is associated with loss of hemispheric lateralization, in the current study we measured the effect of aging on the assumed light source direction. Older participants exhibited, on average, a decreased left bias compared to young participants, as well as greater within-group variability in the distribution of assumed light source directions. In a separate sample of young and old participants, we replicated the age related effect in the assumed light source direction. Furthermore, in both young and old participants the assumed light source direction and the lateralized bias in a line bisection task were correlated. These findings suggest that diminished hemispheric lateralization, which accompanies aging, may affect the perception of the 3D structure of shaded surfaces. Shape from shading may thus provide a simple behavioral tool to track age related changes in hemispheric organization.

Spatial attention can be biased towards an expected dimension.

A commonly held view in both exogenous and endogenous orienting is that spatial attention is associated with enhanced processing of all stimuli at the attended location. However, we often search for a specific target at a particular location, so an observer should be able to jointly specify the target identity and expected location. Whether attention can bias dimension-specific processes at a particular location is not yet clear. We used a dual task to examine the effects of endogenous spatial cues on the accuracy of perceptual judgments of different dimensions. Participants responded to a motion target and a colour target, presented at the same or different locations. We manipulated a central cue to predict the location of the motion or colour target. While overall performance in the two tasks was comparable, cueing effects were larger for the target whose location was predicted by the cue, implying that when attending a particular location, processing of the likely dimension was preferentially enhanced. Additionally, an asymmetry between the motion and colour tasks was seen; motion was modulated by attention, and colour was not. We conclude that attention has some ability to select a dimension at a particular location, indicating integration of spatial and feature-based attention.

Attention to multiple locations is limited by spatial working memory capacity.

What limits the ability to attend several locations simultaneously? There are two possibilities: Either attention cannot be divided without incurring a cost, or spatial memory is limited and observers forget which locations to monitor. We compared motion discrimination when attention was directed to one or multiple locations by briefly presented central cues. The cues were matched for the amount of spatial information they provided. Several random dot kinematograms (RDKs) followed the spatial cues; one of them contained task-relevant, coherent motion. When four RDKs were presented, discrimination accuracy was identical when one and two locations were indicated by equally informative cues. However, when six RDKs were presented, discrimination accuracy was higher following one rather than multiple location cues. We examined whether memory of the cued locations was diminished under these conditions. Recall of the cued locations was tested when participants attended the cued locations and when they did not attend the cued locations. Recall was inaccurate only when the cued locations were attended. Finally, visually marking the cued locations, following one and multiple location cues, equalized discrimination performance, suggesting that participants could attend multiple locations when they did not have to remember which ones to attend. We conclude that endogenously dividing attention between multiple locations is limited by inaccurate recall of the attended locations and that attention poses separate demands on the same central processes used to remember spatial information, even when the locations attended and those held in memory are the same.

Inhibition of return affects contrast sensitivity.

Inhibition of return (IOR)-a slow response to targets at recently attended locations, is believed to play an important role in guiding behaviour. In the attention literature it has been shown that attentional capture by an exogenous cue affects contrast sensitivity so that it alters the appearance of low-contrast stimuli. Despite a significant amount of work over the last quarter century on IOR, it is not yet clear whether IOR operates in the same way. In the current study we examined the effect of IOR on contrast sensitivity-a very early, low-level perceptual process. We found in both a detection task and an orientation discrimination task that lower contrast was needed to detect the stimulus (Experiment 1) and determine its orientation (Experiment 2) at the cued location than at the uncued location, at short cue-target delays, while higher contrast was needed at long delays-reflecting IOR. These results clearly demonstrate that IOR affects contrast sensitivity in a similar way as attentional capture does and suggest that IOR increases perceived contrast of an object in the uncued location.

Cross-cultural effects on the assumed light source direction: evidence from English and Hebrew readers.

When judging the 3D shape of a shaded image, observers generally assume that the light source is placed above and to the left. This leftward bias has been attributed to experiential factors shaped by the observers' handedness or hemispheric dominance. Others have found that experiential factors can rapidly modify the direction of the assumed light source, suggesting a role for learning in shaping perceptual expectations. In the current study, instead, we assessed the contribution of cultural factors affecting the way visual scenes are customarily inspected, in determining the assumed light source direction. Left- and right-handed first language English and Hebrew participants, who read and write from left to right and from right to left, respectively, judged the relative depth of the central hexagon surrounded by six shaded hexagons. We found a left bias in first language English participants, but a significantly smaller one in Hebrew participants. In neither group was the light direction affected by participants' handedness. We conclude that the bias in the assumed light source direction is affected by cultural factors, likely related to the habitual scanning direction employed by participants when reading and writing their first language script.

Matching cue size and task properties in exogenous attention.

Exogenous attention is an involuntary, reflexive orienting response that results in enhanced processing at the attended location. The standard view is that this enhancement generalizes across visual properties of a stimulus. We test whether the size of an exogenous cue sets the attentional field and whether this leads to different effects on stimuli with different visual properties. In a dual task with a random-dot kinematogram (RDK) in each quadrant of the screen, participants discriminated the direction of moving dots in one RDK and localized one red dot. Precues were uninformative and consisted of either a large or a small luminance-change frame. The motion discrimination task showed attentional effects following both large and small exogenous cues. The red dot probe localization task showed attentional effects following a small cue, but not a large cue. Two additional experiments showed that the different effects on localization were not due to reduced spatial uncertainty or suppression of RDK dots in the surround. These results indicate that the effects of exogenous attention depend on the size of the cue and the properties of the task, suggesting the involvement of receptive fields with different sizes in different tasks. These attentional effects are likely to be driven by bottom-up mechanisms in early visual areas.

Hemispheric asymmetry in the remapping and maintenance of visual saliency maps: a TMS study.

Parietal cortex has been implicated in the updating, after eye movements, of a salience map that is required for coherent visual experience and for the control of visually guided behavior. The current experiment investigated whether TMS over anterior intraparietal cortex (AIPCx), just after a saccade, would affect the ability to update and maintain a salience map. In order to generate a salience map, we employed a paradigm in which an uninformative cue was presented at one object in a display to generate inhibition of return (IOR)-an inhibitory tag that renders the cued object less salient than others in the display, and that slows subsequent responses to visual transients at its location. Following the cue, participants made a saccade to either left or right, and we then probed for updating of the location of IOR by measuring manual reaction time to targets appearing at cued location of the cued compared to an uncued object. Between the time of saccade initiation and target appearance, dual-pulse TMS was targeted over right (Experiment 1) or left AIPCx (Experiment 2), and a vertex control side. Updating of the location of IOR was eliminated by TMS over right, but not the left, AIPCx, suggesting that right parietal cortex is involved in the remapping of IOR. Remapping was eliminated by right AIPCx, regardless of whether the saccade was made to the left (contralateral), or right (ipsilateral) visual field, and regardless of which field the target appeared in. We conclude that right AIPCx is the neural substrate for maintaining a salience map across saccades, and not simply for propagating an efference copy of saccade commands.

Is the posner reaction time test more accurate than clinical tests in detecting left neglect in acute and chronic stroke?

UNLABELLED: Rengachary J, d'Avossa G, Sapir A, Shulman GL, Corbetta M. Is the Posner Reaction Time Test more accurate than clinical tests in detecting left neglect in acute and chronic stroke? OBJECTIVE: To compare the accuracy of common clinical tests for left neglect with that of a computerized reaction time Posner test in a stroke population. DESIGN: Neglect measures were collected longitudinally in patients with stroke at the acute ( approximately 2wk) and chronic ( approximately 9mo) stages. Identical measures were collected in a healthy control group. SETTING: Inpatient and outpatient rehabilitation. PARTICIPANTS: Patients with acute stroke (n=59) with left neglect, 30 of whom were tested longitudinally; healthy age-matched controls (n=30). INTERVENTIONS: Not applicable. MAIN OUTCOME MEASURES: A receiver operating characteristic analysis ranking the measures' sensitivity and specificity using a single summary statistic. RESULTS: Most clinical tests were adequately accurate at the acute stage, but many were near chance at the chronic stage. The Posner test was the most sensitive test at both stages. The most sensitive variable was the reaction time difference for detecting targets appearing on the left compared with the right side. CONCLUSIONS: Computerized reaction time tests can be used to screen for subtle but potentially clinically relevant left neglect, which may not be detectable by conventional clinical tests, especially at the chronic stage. Such tests may be useful to assess the severity of the patients' deficits and provide more accurate measures of the degree of recovery in clinical trials than established clinical measures.

Neuroleptics reverse attentional effects in schizophrenia patients.

Patients with schizophrenia show attention deficit characterized by a larger validity effect (fast responses to cued than uncued stimuli) in the right visual field than in the left visual field. In addition, schizophrenia patients do not show inhibition of return (IOR--a mechanism that enables efficient visual search), unless attention is summoned back to the center after the peripheral cue. The present study examined the short-term effect of neuroleptic medications on these two components of visual spatial attention in schizophrenia patients. In order to do this we tested schizophrenia patients that were treated with long-acting neuroleptic medication. These patients were treated once a month, which allowed us to test them with either low or high levels of medication. Here we show that neuroleptic medications reverse the attentional hemispheric asymmetry. In the group with a high level of medication fast RTs to cued trials were found in the right visual field, while in the group with a low level of medication the opposite pattern was found - fast RTs to cued trials were found in the left visual field. In addition, level of medication did not influence IOR - regardless of the level of medication, IOR was observed only when attention was summoned back to the center, unlike control group. These finding suggest an imbalance in dopaminergic activity, possibly in subcortical structures such as the basal ganglia. This study also shows a dissociation between the two components of visual orienting of attention and suggests that facilitation and inhibition are independent processes.

Anatomical correlates of directional hypokinesia in patients with hemispatial neglect.

Unilateral spatial neglect (neglect) is a syndrome characterized by perceptual deficits that prevent patients from attending and responding to the side of space and of the body opposite a damaged hemisphere (contralesional side). Neglect also involves motor deficits: patients may be slower to initiate a motor response to targets appearing in the left hemispace, even when using their unaffected arm (directional hypokinesia). Although this impairment is well known, its anatomical correlate has not been established. We tested 52 patients with neglect after right hemisphere stroke, and conducted an anatomical analysis on 29 of them to find the anatomical correlate of directional hypokinesia. We found that patients with directional hypokinesia had a lesion involving the ventral lateral putamen, the claustrum, and the white matter underneath the frontal lobe. Most importantly, none of the patients without directional hypokinesia had a lesion in the same region. The localization of neglect's motor deficits to the basal ganglia establishes interesting homologies with animal data; it also suggests that a relative depletion of dopamine in the nigrostriatal pathway on the same side of the lesion may be an important pathophysiological mechanism potentially amenable to intervention.

Brain signals for spatial attention predict performance in a motion discrimination task.

The reliability of visual perception is thought to reflect the quality of the sensory information. However, we show that subjects' performance can be predicted, trial-by-trial, by neural activity that precedes the onset of a sensory stimulus. Using functional MRI (fMRI), we studied how neural mechanisms that mediate spatial attention affect the accuracy of a motion discrimination judgment. The amplitude of blood oxygen level-dependent (BOLD) signals after a cue directing spatial attention predicted subjects' accuracy on 60-75% of the trials. Widespread predictive signals, which included dorsal parietal, visual extra-striate, prefrontal and sensory-motor cortex, depended on whether the cue correctly specified the stimulus location. Therefore, these signals indicate the degree of utilization of the cued information and play a role in the control of spatial attention. We conclude that variability in perceptual performance can be partly explained by the variability in endogenous, preparatory processes and that BOLD signals can be used to forecast human behavior.

Neural basis and recovery of spatial attention deficits in spatial neglect.

The syndrome of spatial neglect is typically associated with focal injury to the temporoparietal or ventral frontal cortex. This syndrome shows spontaneous partial recovery, but the neural basis of both spatial neglect and its recovery is largely unknown. We show that spatial attention deficits in neglect (rightward bias and reorienting) after right frontal damage correlate with abnormal activation of structurally intact dorsal and ventral parietal regions that mediate related attentional operations in the normal brain. Furthermore, recovery of these attention deficits correlates with the restoration and rebalancing of activity within these regions. These results support a model of recovery based on the re-weighting of activity within a distributed neuronal architecture, and they show that behavioral deficits depend not only on structural changes at the locus of injury, but also on physiological changes in distant but functionally related brain areas.

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.

Parietal lobe lesions disrupt saccadic remapping of inhibitory location tagging.

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 environmentally based reference frame.

Attending to the thalamus: inhibition of return and nasal-temporal asymmetry in the pulvinar.

Inhibition of return (IOR) is a mechanism whereby the attentional system favors novel locations by inhibiting already scanned ones. An important question is what the neural structures are involved. Recently, we studied a patient with damage to the superior colliculus (SC) and concluded that the SC generates IOR. However, it is possible that IOR is generated beyond the colliculus, for example, by the pulvinar. In this paper we tested three patients with unilateral damage to the pulvinar and demonstrated that the pulvinar is not necessary for IOR generation, providing additional support to the suggestion that the SC generates IOR. In addition, since we used monocular presentation, we were able to furnish behavioral evidence for nasal-temporal asymmetrical representation of visual input in the pulvinar.