Atypical attentional filtering of visual information in youth with chromosome 22q11.2 deletion syndrome as indexed by event-related potentials.

BACKGROUND: Youth with chromosome 22q11.2 deletion syndrome (22q) face one of the highest genetic risk factors for the development of schizophrenia. Previous research suggests impairments in attentional control and potential interactions with elevated anxiety and reduced adaptive functioning may increase the risk for developing psychosis in this population. Here, we examined how variations in attentional control relate to the presence or severity of psychosis-proneness symptoms in these individuals. METHODS: To achieve this, we measured attentional control in youth (12-18 years) with 22q (N = 35) compared to a typically developing group (N = 45), using a flanker task (the Distractor Target task) while measuring neural activity with event-related potentials. RESULTS: Similar to previous findings observed in people with schizophrenia, greater attentional capture by, and reduced suppression of, non-target flanker stimuli characterized participants with 22q and was indexed by the N2pc (N2-posterior-contralateral) and PD (distractor positivity) components. Although we observed no relationships between these components and measures of psychosis-proneness in youth with 22q, these individuals endorsed a relatively low incidence of positive symptoms overall. CONCLUSIONS: Our results provide neural evidence of an attentional control impairment in youth with 22q that suggests these individuals experience sustained attentional focus on irrelevant information and reduced suppression of distracting stimuli in their environment. Impairments in attentional control might be a valid biomarker of the potential to develop attenuated positive symptoms or frank psychosis in high-risk individuals long before the age at which such symptoms typically arise. The evaluation of such a hypothesis, and the preventive potential for the putative biomarker, should be the focus of future studies.

Cognitive and neural mechanisms of visual search.

The visual search paradigm is a useful tool for studying the perception of multiple-element stimulus arrays, and recent studies have identified a number of factors that influence search performance. Of special importance is the selection process, and recent neurophysiological experiments have shown that the suppression of information from irrelevant objects plays a crucial role in the selection of visual search targets.

How does attention attenuate target-distractor interference in vision?. Evidence from magnetoencephalographic recordings.

This study used magnetoencephalographic and electroencephalographic recordings to investigate the neural mechanisms that underlie the attentional resolution of ambiguous feature coding in visual search. We addressed this issue by comparing neural activity related to target discrimination under conditions of more versus less feature overlap between the target and distractor items. The results show that increasing feature overlap leads to a focal enhancement of neural activity in ventral occipito-temporal areas, consistent with the larger need to attenuate distractor interference. Furthermore, the results suggest that distractor attenuation proceeds as a stepwise operation, with different spatial locations containing interfering features being suppressed successively. These findings support theories of visual search that emphasize location-based attentional selection as a key mechanism in resolving ambiguous feature coding in vision.

Sequential operation of disconnected cerebral hemispheres in split-brain patients.

When two concurrent sensorimotor tasks require separate responses, selection of the first response generally delays selection of the second. Dual-task performance was examined in four patients who had undergone surgical transection of the forebrain commissures including the corpus callosum. One light flash was presented to each visual field in succession, and patients made a choice response to each stimulus with the ipsilateral hand, thereby confining the tasks to separate hemispheres. All four showed dual-task interference very similar to that found with normal individuals. Therefore, still-intact subcortical structures must play a critical role in sequencing response selection processes (the 'dual-task bottleneck'), confirming the distinction between the attentional limitations involved in planning actions and those involved in perceptual analysis.

Multiple mechanisms of visual-spatial attention: recent evidence from human electrophysiology.

Natural visual scenes contain vast quantities of information--far more than the visual system can process in a short period of time-and spatial attention is therefore used to focus the visual system's processing resources onto a subset of the incoming visual information. Most psychological theories of attention posit a single mechanism for this focusing of attention, but recent electrophysiological studies have provided evidence that the visual system employs several separable neural mechanisms of spatial attention. This paper describes the evidence for multiple attentional mechanisms and suggests links between these neurophysiologically defined mechanisms and specific functional processes that have been proposed in psychological theories of attention.

Spatial filtering during visual search: evidence from human electrophysiology.

The identification of targets in visual search arrays may be improved by suppressing competing information from the surrounding distractor items. The present study provided evidence that this hypothetical filtering process has a neural correlate, the "N2pc" component of the event-related potential waveform. The N2pc was observed when a target item was surrounded by competing distractor items but was absent when the array could be rejected as a nontarget on the basis of simple feature information. In addition, the N2pc was eliminated when filtering was discouraged by removing the distractor items, making the distractors relevant, or making all items within an array identical. Combined with previous topographic analyses, these results suggest that attentional filtering occurs in occipital cortex under the control of feedback from higher cortical regions after a preliminary feature-based analysis of the stimulus array.

Storage of features, conjunctions and objects in visual working memory.

Working memory can be divided into separate subsystems for verbal and visual information. Although the verbal system has been well characterized, the storage capacity of visual working memory has not yet been established for simple features or for conjunctions of features. The authors demonstrate that it is possible to retain information about only 3-4 colors or orientations in visual working memory at one time. Observers are also able to retain both the color and the orientation of 3-4 objects, indicating that visual working memory stores integrated objects rather than individual features. Indeed, objects defined by a conjunction of four features can be retained in working memory just as well as single-feature objects, allowing many individual features to be retained when distributed across a small number of objects. Thus, the capacity of visual working memory must be understood in terms of integrated objects rather than individual features.

Mechanisms of visual-spatial attention: resource allocation or uncertainty reduction?

Many studies have found that stimuli can be discriminated more accurately at attended locations than at unattended locations, and such results have typically been taken as evidence for the hypothesis that attention operates by allocating limited perceptual processing resources to attended locations. An alternative proposal, however, is that attention acts to reduce uncertainty about target location, thereby increasing accuracy by decreasing the number of noise sources. To distinguish between these alternatives, we conducted 6 spatial cuing experiments in which target location uncertainty was eliminated. Despite the absence of uncertainty, target discriminations were more accurate at the attended location, consistent with resource allocation models. These cue validity effects were observed under a broad range of conditions, including central and peripheral cuing, but were absent at very short cue-target delay intervals.

Electrophysiological evidence for a postperceptual locus of suppression during the attentional blink.

When an observer detects a target in a rapid stream of visual stimuli, there is a brief period of time during which the detection of subsequent targets is impaired. In this study, event-related potentials (ERPs) were recorded from normal adult observers to determine whether this "attentional blink" reflects a suppression of perceptual processes or an impairment in postperceptual processes. No suppression was observed during the attentional blink interval for ERP components corresponding to sensory processing (the P1 and N1 components) or semantic analysis (the N400 component). However, complete suppression was observed for an ERP component that has been hypothesized to reflect the updating of working memory (the P3 component). Results indicate that the attentional blink reflects an impairment in a postperceptual stage of processing.

Visual attention modulates signal detectability.

The mechanism by which visual-spatial attention affects the detection of faint signals has been the subject of considerable debate. It is well known that spatial cuing speeds signal detection. This may imply that attentional cuing modulates the processing of sensory information during detection or, alternatively, that cuing acts to create decision bias favoring input at the cued location. These possibilities were evaluated in 3 spatial cuing experiments. Peripheral cues were used in Experiment 1 and central cues were used in Experiments 2 and 3. Cuing similarly enhanced measured sensitivity, P(A) and d', for simple luminance detection in all 3 experiments. Under some conditions it also induced shifts in decision criteria (beta). These findings indicate that visual-spatial attention facilitates the processing of sensory input during detection either by increasing sensory gain for inputs at cued locations or by prioritizing the processing of cued inputs.

Effects of spatial cuing on luminance detectability: psychophysical and electrophysiological evidence for early selection.

Three experiments were conducted to determine whether attention-related changes in luminance detectability reflect a modulation of early sensory processing. Experiments 1 and 2 used peripheral cues to direct attention and found substantial effects of cue validity on target detectability; these effects were consistent with a sensory-level locus of selection but not with certain memory- or decision-level mechanisms. In Experiment 3, event-related brain potentials were recorded in a similar paradigm using central cues, and attention was found to produce changes in sensory-evoked brain activity beginning within the 1st 100 ms of stimulus processing. These changes included both an enhancement of sensory responses to attended stimuli and a suppression of sensory responses to unattended stimuli; the enhancement and suppression effects were isolated to different neural responses, indicating that they may arise from independent attentional mechanisms.

Are attentional dwell times inconsistent with serial visual search?

Duncan, Ward, and Shapiro (1994) estimated that attention must remain focused on an object for several hundred milliseconds before being shifted to another object, and they referred to this period as theattentional dwell time. An important implication of these long estimates of the dwell time for models of visual search is that the search process must not involve an item-by-item serial scanning mechanism. If it did, then searching through an array of items would require enormous amounts of time, which-based on data from visual search experiments-it does not. The present report, however, provides evidence that the long estimates of attentional dwell time were caused, at least in part, by the use of masked targets. Implications of these variable estimates of the attentional dwell time for models of visual search are discussed.

Direct neurophysiological evidence for spatial suppression surrounding the focus of attention in vision.

The spatial focus of attention has traditionally been envisioned as a simple spatial gradient of enhanced activity that falls off monotonically with increasing distance. Here, we show with high-density magnetoencephalographic recordings in human observers that the focus of attention is not a simple monotonic gradient but instead contains an excitatory peak surrounded by a narrow inhibitory region. To demonstrate this center-surround profile, we asked subjects to focus attention onto a color pop-out target and then presented probe stimuli at various distances from the target. We observed that the electromagnetic response to the probe was enhanced when the probe was presented at the location of the target, but the probe response was suppressed in a narrow zone surrounding the target and then recovered at more distant locations. Withdrawing attention from the pop-out target by engaging observers in a demanding foveal task eliminated this pattern, confirming a truly attention-driven effect. These results indicate that neural enhancement and suppression coexist in a spatially structured manner that is optimal to attenuate the most deleterious noise during visual object identification.

Direct and indirect integration of event-related potentials, functional magnetic resonance images, and single-unit recordings.

Cognitive neuroimaging techniques vary along three primary dimensions: invasiveness, temporal resolution, and spatial resolution. Several of the major techniques excel on two of these three dimensions, but none of them excels on all three. In principle, multiple techniques with different strengths and weaknesses could be combined to obtain high temporal and spatial resolution data about human neural activity, and this article compares two approaches to combining microelectrode, hemodynamic, and electromagnetic measures of neural activity. The first approach involves using structural magnetic resonance images to provide a common reference frame for the mathematical estimation of neural activity, and the second approach involves parallel experimental manipulations and converging evidence. At present, neither approach is entirely satisfactory, and the integration of different measures of neural activity, therefore, requires a combination of direct and indirect approaches.

Electrophysiological evidence for parallel and serial processing during visual search.

Event-related potentials were recorded from young adults during a visual search task in order to evaluate parallel and serial models of visual processing in the context of Treisman's feature integration theory. Parallel and serial search strategies were produced by the use of feature-present and feature-absent targets, respectively. In the feature-absent condition, the slopes of the functions relating reaction time and latency of the P3 component to set size were essentially identical, indicating that the longer reaction times observed for larger set sizes can be accounted for solely by changes in stimulus identification and classification time, rather than changes in post-perceptual processing stages. In addition, the amplitude of the P3 wave on target-present trials in this condition increased with set size and was greater when the preceding trial contained a target, whereas P3 activity was minimal on target-absent trials. These effects are consistent with the serial self-terminating search model and appear to contradict parallel processing accounts of attention-demanding visual search performance, at least for a subset of search paradigms. Differences in ERP scalp distributions further suggested that different physiological processes are utilized for the detection of feature presence and absence.

Electrophysiological correlates of feature analysis during visual search.

Event-related brain potentials (ERPs) were recorded from normal young adults during visual search tasks in which the stimulus arrays contained either eight identical items (homogeneous arrays) or seven identical items and one deviant item (pop-out arrays). Four experiments were conducted in which different classes of stimulus arrays were designated targets and the remaining stimulus arrays were designated nontargets. In Experiments 1 and 2, both target and nontarget pop-out stimuli elicited an enhanced anterior N2 wave and a contralaterally larger posterior P1 wave, but Experiments 3 and 4 demonstrated that these components do not reflect fully automatic pop-out detection processes. In all four experiments, target pop-outs elicited enlarged anterior P2, posterior N2, occipital P3, and parietal P3 waves. The target-elicited posterior N2 wave contained a contralateral subcomponent (N2pc) that exhibited a focus over occipital cortex in maps of current source density. The overall pattern of results was consistent with guided search models in which preattentive stimulus information is used to guide attention to task-relevant stimuli.

The capacity of visual working memory for features and conjunctions.

Short-term memory storage can be divided into separate subsystems for verbal information and visual information, and recent studies have begun to delineate the neural substrates of these working-memory systems. Although the verbal storage system has been well characterized, the storage capacity of visual working memory has not yet been established for simple, suprathreshold features or for conjunctions of features. Here we demonstrate that it is possible to retain information about only four colours or orientations in visual working memory at one time. However, it is also possible to retain both the colour and the orientation of four objects, indicating that visual working memory stores integrated objects rather than individual features. Indeed, objects defined by a conjunction of four features can be retained in working memory just as well as single-feature objects, allowing sixteen individual features to be retained when distributed across four objects. Thus, the capacity of visual working memory must be understood in terms of integrated objects rather than individual features, which places significant constraints on cognitive and neurobiological models of the temporary storage of visual information.

The visual N1 component as an index of a discrimination process.

Many previous studies have demonstrated that the visual N1 component is larger for attended-location stimuli than for unattended-location stimuli. This difference is observed typically only for tasks involving a discrimination of the attended-location stimuli, suggesting that the N1 wave reflects a discrimination process that is applied to the attended location. The present study tested this hypothesis by examining the N1 component elicited by attended stimuli under conditions that either required or did not require the subject to perform a discrimination. Specifically, the N1 elicited by foveal stimuli during choice-reaction time (RT) tasks was compared with the N1 elicited by identical stimuli during simple-RT tasks. In three experiments, a larger posterior N1 was observed in choice-RT tasks than in simple-RT tasks, even when several potential confounds were eliminated (e.g., arousal and motor preparation). This N1 discrimination effect was observed even when no motor response was required and was present for both color- and form-based discriminations. Moreover, this discrimination effect was equally large for easy and difficult discriminations, arguing against a simple resource-based explanation of the present results. Instead, the results of this study are consistent with the hypothesis that the visual N1 component reflects the operation of a discrimination process within the focus of attention.

The role of attention in feature detection and conjunction discrimination: an electrophysiological analysis.

An electrophysiological probe technique was used to ascertain whether the same attentional mechanisms are employed for both the detection of simple visual features and the discrimination of conjunctions of features. Visual search arrays containing 14 grey items and 2 colored items were presented; one color was designated relevant for each trial block. Subjects were required to report the presence or absence of the relevant color (feature detection condition) or the shape of the relevantly-colored item (conjunction discrimination condition). Shortly after the onset of the search array, a task-irrelevant probe stimulus was flashed at the location of the relevant or irrelevant color and the event-related potential (ERP) produced by this stimulus was used to assess sensory processing at the probed location. Probes presented at the location of the relevant color were found to elicit enhanced ERP components and probes presented on the opposite side of the display from the relevant color elicited suppressed components. These effects were observed in both the detection and discrimination conditions, indicating that spatially restricted attentional processes are used for both the detection of simple features and the discrimination of conjunctions. However, one ERP component (the PI wave) exhibited these effects in the discrimination condition but not in the detection condition, indicating that conjunction discrimination utilizes additional attentional processes beyond those required for feature detection.