Spatial probability cuing and right hemisphere damage

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

Poor hand-pointing to sounds in right brain-damaged patients: Not just a problem of spatial-hearing

We asked 22 right brain-damaged (RBD) patients and 11 elderly healthy controls to perform hand-pointing movements to free-field unseen sounds, while modulating two non-auditory variables: the initial position of the responding hand (left, centre or right) and the presence or absence of task-irrelevant ambient vision. RBD patients suffering from visual neglect, unlike RBD patients without neglect and healthy controls, showed a systematic rightward error in sound localisation, which was modulated by the non-auditory variables. Localisation errors were exacerbated by initial hand-position to the right of the body-midline, and reduced by the leftwards initial hand-position. Moreover, for the visual neglect patients, mere presence of ambient vision worsened localisation errors. These results demonstrate that although hand-pointing to sounds has often been considered a straightforward approach to investigate sound-localisation abilities in brain-damaged patients, in some patients it may actually reveal localisation deficits that reflect a combination of impaired spatial-hearing and spatial biases from other sensory modalities (i.e., vision and proprioception).     

Poor hand-pointing to sounds in right brain-damaged patients: not just a problem of spatial-hearing

We asked 22 right brain-damaged (RBD) patients and 11 elderly healthy controls to perform hand-pointing movements to free-field unseen sounds, while modulating two non-auditory variables: the initial position of the responding hand (left, centre or right) and the presence or absence of task-irrelevant ambient vision. RBD patients suffering from visual neglect, unlike RBD patients without neglect and healthy controls, showed a systematic rightward error in sound localisation, which was modulated by the non-auditory variables. Localisation errors were exacerbated by initial hand-position to the right of the body-midline, and reduced by the leftwards initial hand-position. Moreover, for the visual neglect patients, mere presence of ambient vision worsened localisation errors. These results demonstrate that although hand-pointing to sounds has often been considered a straightforward approach to investigate sound-localisation abilities in brain-damaged patients, in some patients it may actually reveal localisation deficits that reflect a combination of impaired spatial-hearing and spatial biases from other sensory modalities (i.e., vision and proprioception).     

Clock drawing in spatial neglect: A comprehensive analysis of clock perimeter, placement, and accuracy

Clock drawings produced by right-brain-damaged (RBD) individuals with spatial neglect often contain an abundance of empty space on the left while numbers and hands are placed on the right. However, the clock perimeter is rarely compromised in neglect patients' drawings. By analysing clock drawings produced by 71 RBD and 40 healthy adults, this study investigated whether the geometric characteristics of the clock perimeter reveal novel insights to understanding spatial neglect. Neglect participants drew smaller clocks than either healthy or non-neglect RBD participants. While healthy participants' clock perimeter was close to circular, RBD participants drew radially extended ellipses. The mechanisms for these phenomena were investigated by examining the relation between clock-drawing characteristics and performance on six subtests of the Behavioral Inattention Test (BIT). The findings indicated that the clock shape was independent of any BIT subtest or the drawing placement on the test sheet and that the clock size was significantly predicted by one BIT subtest: the poorer the figure and shape copying, the smaller the clock perimeter. Further analyses revealed that in all participants, clocks decreased in size as they were placed farther from the centre of the paper. However, even when neglect participants placed their clocks towards the centre of the page, they were smaller than those produced by healthy or non-neglect RBD participants. These results suggest a neglect-specific reduction in the subjectively available workspace for graphic production from memory, consistent with the hypothesis that neglect patients are impaired in the ability to enlarge the attentional aperture.     

Spatial probability as an attentional cue in visual search

We investigated the role of spatial probabilities in target location during participants' performance of a visual search task. Experiments 1 and 2 demonstrated that spatial probabilities could serve as a powerful attentional bias that produced faster detection of targets in high-probability locations than of those in low- or random-probability locations. The effect could not be explained by repetition priming alone. Moreover, responses to targets in low-probability locations were slowed only when a distractor was present in the high-probability location. In Experiments 3-5, we compared the effects of spatial probability with an explicit endogenous cue and a salient exogenous cue. Facilitation due to spatial probability was independent of any benefit afforded by the explicit endogenous cue but interacted with the salient exogenous cue, such that the exogenous cue validity effect was compressed for targets in the high-probability location. Together, these results suggest that the spatial probabilities governing target location constitute a potent bias of visual processing and, as such, can be considered an attentional cue that differs from both typical explicit endogenous and salient exogenous cues.     

A consistent but non-coincident visual pattern facilitates the learning of spatial relations among locations

Human participants searched in a dynamic three-dimensional computer-generated virtual-environment open-field search task for four hidden goal locations arranged in a diamond configuration located in a 5 × 5 matrix of raised bins. Participants were randomly assigned to one of two groups: visual pattern or visual random. All participants experienced 30 trials in which four goal locations maintained the same spatial relations to each other (i.e., a diamond pattern), but this diamond pattern moved to random locations within the 5 × 5 matrix from trial to trial. For participants in the visual pattern group, four locations were marked in a distinct color and arranged in a diamond pattern that moved to a random location independent of the hidden spatial pattern from trial to trial throughout the experimental session. For participants in the visual random group, four random locations were marked with a distinct color and moved to random locations independent from the hidden spatial pattern from trial to trial throughout the experimental session. As a result, the visual cues for the visual pattern group were consistent but not coincident with the hidden spatial pattern, whereas the visual cues for the visual random group were neither consistent nor coincident with the hidden spatial pattern. Results indicated that participants in both groups learned the spatial configuration of goal locations and that the presence of consistent but noncoincident visual cues facilitated the learning of spatial relations among locations.     

Short-term and long-term attentional biases to frequently encountered target features

It has long been known that frequently occurring targets are attended better than infrequent ones in visual search. But does this frequency-based attentional prioritization reflect momentary or durable changes in attention? Here we observed both short-term and long-term attentional biases for visual features as a function of different types of statistical associations between the targets, distractors, and features. Participants searched for a target, a line oriented horizontally or vertically among diagonal distractors, and reported its length. In one set of experiments we manipulated the target’s color probability: Targets were more often in Color 1 than in Color 2. The distractors were in other colors. Participants found Color 1 targets more quickly than Color 2 targets, but this preference disappeared immediately when the target’s color became random in the subsequent testing phase. In the other set of experiments, we manipulated the diagnostic values of the two colors: Color 1 was more often a target than a distractor; Color 2 was more often a distractor than a target. Participants found Color 1 targets more quickly than Color 2 targets. Importantly, and in contrast to the first set of experiments, the featural preference was sustained in the testing phase. These results suggest that short-term and long-term attentional biases are products of different statistical information. Finding a target momentarily activates its features, inducing short-term repetition priming. Long-term changes in attention, on the other hand, may rely on learning diagnostic features of the targets.     

Implicit short- and long-term memory direct our gaze in visual search

Visual attention is strongly affected by the past: both by recent experience and by long-term regularities in the environment that are encoded in and retrieved from memory. In visual search, intertrial repetition of targets causes speeded response times (short-term priming). Similarly, targets that are presented more often than others may facilitate search, even long after it is no longer present (long-term priming). In this study, we investigate whether such short-term priming and long-term priming depend on dissociable mechanisms. By recording eye movements while participants searched for one of two conjunction targets, we explored at what stages of visual search different forms of priming manifest. We found both long- and short- term priming effects. Long-term priming persisted long after the bias was present, and was again found even in participants who were unaware of a color bias. Short- and long-term priming affected the same stage of the task; both biased eye movements towards targets with the primed color, already starting with the first eye movement. Neither form of priming affected the response phase of a trial, but response repetition did. The results strongly suggest that both long- and short-term memory can implicitly modulate feedforward visual processing.     

Visual attention amplifies response priming of pointing movements to color targets

We studied the influence of spatial visual attention on the time course of primed pointing movements. We measured pointing responses to color targets preceded by color stimuli priming either the same response as the target or the opposite response. The effects of visual attention at the prime and target locations were studied by varying both the cue-prime and prime-target intervals when presenting either exogenous attentional cues or, in a separate experiment, endogenous cues whose processing was a precondition for performing the task. Pointing trajectories revealed large priming effects in which pointing responses were first controlled by prime signals and then captured in midflight by target signals. Priming effects were strongly amplified when the relevant prime locations were visually attended at optimal cue-prime SOAs, with attention modulating the entire time course of the primed pointing movements. We propose that visual attention amplifies the earliest waves of visuomotor feedforward information, elicited in turn by primes and by targets.     

Probability cuing of target location facilitates visual search implicitly in normal participants and patients with hemispatial neglect

We explored how variability in the probability of target locations affects visual search in normal individuals and in patients with hemispatial neglect, a deficit in attending to the contralesional side of space. Young and elderly normal participants responded faster when targets appeared in the more probable region than when targets appeared in the less probable region. Similarly, patients were sensitive to the distribution of targets, even in the neglected field. Although the attentional gradient that characterizes neglect was not eliminated, the response facilitation due to the probability distribution was proportionate to that of control participants and equal in magnitude across the neglected field. All participants exploited the uneven distribution of targets to enhance task performance without explicit instructions to do so or awareness of biases in their behavior. These results suggest that attentional orientation and sensitivity to external probabilities are possibly dissociable. An early sensory and a late motor mechanism are postulated as possibly being involved in the observed probability–matching behavior of participants.     

Processing of distractors inside and outside the attentional focus in a priming procedure

A priming procedure was used to study the processing of distractors located either inside (between the location of two targets) or outside (peripherally to the locations of the targets) the focus of attention. The stimuli were five-letter arrays, and participants had to decide whether two marked target letters were the same or different. In Experiments 1 and 2, positive priming was obtained both when targets and in-distractors in primes repeated as targets in probes; negative priming was found when out-distractor primes repeated as targets in probes. In Experiment 3, we also manipulated the match in letter case from primes to probes. In-distractors produced reliable positive priming, irrespective of whether the letters matched in case. In contrast out-distractors produced negative priming but only when the letters had the same case in primes and probes. These results are attributed to a spatial attention process operating (in this case) on low-level visual features, and an object-based selection process that enables more abstract information to be processed for selected stimuli.     

Acquisition of habitual visual attention and transfer to related tasks

Extensive research has shown that statistical learning affects perception, attention, and action control; however, few studies have directly linked statistical learning with the formation of habits. Evidence that learning can induce a search habit has come from location probability learning, in which people prioritize locations frequently attended to in the past. Here, using an alternating training–testing procedure, we demonstrated that the initial attentional bias arises from short-term intertrial priming, whereas probability learning takes longer to emerge, first reaching significance in covert orienting (measured by reaction times) after about 48 training trials, and in overt orienting (measured by eye movements) after about 96 training trials. We further showed that location probability learning is persistent after training is discontinued, by transferring from a letter search task to a scene search task—emulating another characteristic feature of habits. By identifying the onset of probability learning and investigating its task specificity, this study provides evidence that probability cuing can induce habitual spatial attention.     

All set! Evidence of simultaneous attentional control settings for multiple target colors

Although models of visual search have often assumed that attention can only be set for a single feature or property at a time, recent studies have suggested that it may be possible to maintain more than one attentional control setting. The aim of the present study was to investigate whether spatial attention could be guided by multiple attentional control settings for color. In a standard spatial cueing task, participants searched for either of two colored targets accompanied by an irrelevantly colored distractor. Across five experiments, results consistently showed that nonpredictive cues matching either target color produced a significant spatial cueing effect, while irrelevantly colored cues did not. This was the case even when the target colors could not be linearly separated from irrelevantly cue colors in color space, suggesting that participants were not simply adopting one general color set that included both target colors. The results could not be explained by intertrial priming by previous targets, nor could they be explained by a single inhibitory set for the distractor color. Overall, the results are most consistent with the maintenance of multiple attentional control settings.     

Experience-guided attention: Uniform and implicit

Frequently finding a visual search target in one region of space induces a spatial attentional bias toward that region. Past studies on this effect typically tested fewer than 20 participants. The small sample prevents an investigation of two properties of learning: visual field uniformity and role of explicit awareness. Pooling data from multiple studies, here we examined location probability learning from ~120,000 visual search trials across 420 participants. Participants performed a serial search task. Unbeknownst to them, the target was disproportionately likely to appear in one visual quadrant. Location probability learning (LPL) was measured as the difference in reaction time to targets in the high-probability “rich” quadrant and the low-probability “sparse” quadrants. Results showed a lack of visual field effect. LPL was equivalent for “rich” quadrant in the upper left, upper right, lower left, and lower right. Learning did not induce a hotspot diagonal to the “rich” quadrant. To the contrary, RT was the longest in the diagonal quadrant. Recognition rate of the “rich” quadrant was above chance. However, recognition accuracy was unrelated to the size of LPL. Implicit learning induces visual-field-independent changes in spatial attention.     

Distinct,but top-down modulable color and positional priming mechanisms in visual pop-out search

Three experiments examined reaction time (RT) performance in visual pop-out search. Search displays comprised of one color target and two distractors which were presented at 24 possible locations on a circular ellipse. Experiment 1 showed that re-presentation of the target at a previous target location led to expedited RTs, whereas presentation of the target at a distractor location led to slowed RTs (relative to target presentation at a previous empty location). RTs were also faster when the color of the target was the same across consecutive trials, relative to a change of the target’s color. This color priming was independent of the positional priming. Experiment 2 revealed larger positional facilitation, relative to Experiment 1, when position repetitions occurred more likely than chance level; analogously, Experiment 3 revealed stronger color priming effects when target color repetitions were more likely. These position and color manipulations did not change the pattern of color (Experiment 2) and positional priming effects (Experiment 3). While these results support the independency of color and positional priming effects (e.g., Maljkovic and Nakayama in Percept Psychophys 58:977–991, 1996), they also show that these (largely ‘automatic’) effects are top-down modulable when target position and color are predictable (e.g., Müller et al. in Vis Cogn 11:577–602, 2004).     

Hemispatial neglect and serial order in verbal working memory

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

Probing Representations of Gymnastics Movements: A Visual Priming Study

In this study, we designed a visual short‐term priming paradigm to investigate the mechanisms underlying the priming of movements and to probe movement representations in motor experts and matched controls. We employed static visual stimuli that implied or not human whole‐body movements, that is, gymnastics movements and static positions. Twelve elite female gymnasts and twelve matched controls performed a speeded two‐choice response time task. The participants were presented with congruent and incongruent prime‐target pairs and had to decide whether the target stimulus represented a gymnastics movement or a static position. First, a visual priming effect was observed in the two groups. Second, a stimulus–response rote association could not easily account for our results. Novel primes never presented as targets could also prime the targets. Third, by manipulating three levels of prime‐target relations in moving congruent pairs, we demonstrated that the more similar prime‐target pairs, the greater the facilitation in target. Lastly, gymnastics motor expertise impacted on priming effects.     

On the role of attention in generating explicit awareness of contingent relations: evidence from spatial priming

In a series of four experiments, we examine the hypothesis that selective attention is crucial for the generation of conscious knowledge of contingency information. We investigated this question using a spatial priming task in which participants were required to localize a target letter in a probe display. In Experiment 1, participants kept track of the frequency with which the predictive letter in the prime appeared in various locations. This manipulation had a negligible impact on contingency awareness. Subsequent experiments requiring participants to attend to features (color, location) of the predictive letter increased contingency awareness somewhat, but there remained a large proportion of individuals who remained unaware of the strong contingency. Together the results of our experiments suggest that the construct of attention does not fully capture the processes that lead to contingency awareness, and suggest a critical role for bottom-up feature integration in explicit contingency learning.     

Kinematics of Goal-Directed Arm Movements in Neglect: Control of Hand Velocity

Do patients with unilateral neglect exhibit direction-specific deficits in the control of movement velocity when performing goal-directed arm movements? Five patients with left-sided neglect performed unrestrained three-dimensional pointing movements to visual targets presented at body midline, the left and right hemispace. A group of healthy adults and a group of patients with right-hemispheric brain damage but no neglect served as controls. Pointing was performed under normal room light or in darkness. Time-position data of the hand were recorded with an opto-electronic camera system. We found that compared to healthy controls, movement times were longer in both patient groups due to prolonged acceleration and deceleration phases. Tangential peak hand velocity was lower in both patient groups, but not significantly different from controls. Single peak, bell-shaped velocity profiles of the hand were preserved in all right hemispheric patients and in three out of five neglect patients. Most important, the velocity profiles of neglect patients to leftward targets did not differ significantly from those to targets in the right hemispace. In summary, we found evidence for general bradykinesia in neglect patients, but not for a direction-specific deficit in the control of hand velocity. We conclude that visual neglect induces characteristic changes in exploratory behavior, but not in the kinematics of goal-directed movements to objects in peripersonal space.     

Facilitation of learning spatial relations among locations by visual cues: Implications for theoretical accounts of spatial learning

Human participants searched in a real environment or interactive 3-D virtual environment open field for four hidden goal locations arranged in a 2 × 2 square configuration in a 5 × 5 matrix of raised bins. The participants were randomly assigned to one of two groups: cues 1 pattern or pattern only. The participants experienced a training phase, followed by a testing phase. Visual cues specified the goal locations during training only for the cues 1 pattern group. Both groups were then tested in the absence of visual cues. The results in both environments indicated that the participants learned the spatial relations among goal locations. However, visual cues during training facilitated learning of the spatial relations among goal locations: In both environments, the participants trained with the visual cues made fewer errors during testing than did those trained only with the pattern. The results suggest that learning based on the spatial relations among locations may not be susceptible to cue competition effects and have implications for standard associative and dual-system accounts of spatial learning.