Isolating the Neural Substrates of Visually Guided Attention Orienting in Humans.

The neural processes that enable healthy humans to orient attention to sudden visual events are poorly understood because they are tightly intertwined with purely sensory processes. Here we isolated visually guided orienting activity from sensory activity using event-related potentials (ERPs). By recording ERPs to a lateral stimulus and comparing waveforms obtained under conditions of attention and inattention, we identified an early positive deflection over the ipsilateral visual cortex that was associated with the covert orienting of visual attention to the stimulus. Across five experiments with male and female adult participants, this ipsilateral visual orienting activity (VOA) could be distinguished from purely sensory-evoked activity and from other top-down spatial attention effects. The VOA was linked with behavioral measures of orienting, being significantly larger when the stimulus was detected rapidly than when it was detected more slowly, and its presence was independent of saccadic eye movements toward the targets. The VOA appears to be a specific neural index of the visually guided orienting of attention to a stimulus that appears abruptly in an otherwise uncluttered visual field.SIGNIFICANCE STATEMENT The study of visual attention orienting has been an important impetus for the field of cognitive neuroscience. Seminal reaction-time studies demonstrated that a suddenly appearing visual stimulus attracts attention involuntarily, but the neural processes associated with visually guided attention orienting have been difficult to isolate because they are intertwined with sensory processes that trigger the orienting. Here, we disentangled orienting activity from sensory activity using scalp recordings of event-related electrical activity in the human brain. A specific neural index of visually guided attention orienting was identified. Surprisingly, whereas peripheral sensory stimulation is processed initially and predominantly by the contralateral visual cortex, this electrophysiological index of visual orienting was recorded over the cerebral hemisphere that was ipsilateral to the attention-capturing stimulus.

Inability to suppress salient distractors predicts low visual working memory capacity.

According to contemporary accounts of visual working memory (vWM), the ability to efficiently filter relevant from irrelevant information contributes to an individual's overall vWM capacity. Although there is mounting evidence for this hypothesis, very little is known about the precise filtering mechanism responsible for controlling access to vWM and for differentiating low- and high-capacity individuals. Theoretically, the inefficient filtering observed in low-capacity individuals might be specifically linked to problems enhancing relevant items, suppressing irrelevant items, or both. To find out, we recorded neurophysiological activity associated with attentional selection and active suppression during a competitive visual search task. We show that high-capacity individuals actively suppress salient distractors, whereas low-capacity individuals are unable to suppress salient distractors in time to prevent those items from capturing attention. These results demonstrate that individual differences in vWM capacity are associated with the timing of a specific attentional control operation that suppresses processing of salient but irrelevant visual objects and restricts their access to higher stages of visual processing.

Mental rotation requires visual short-term memory: evidence from human electric cortical activity.

The purpose of the present study was to seek evidence that mental rotation is accomplished by transforming a representation held in visual short-term memory (VSTM). In order to accomplish this goal, we utilized the sustained posterior contralateral negativity (SPCN), an electrophysiological index of the maintenance of information in VSTM. We hypothesized that if mental rotation is accomplished by transforming a representation held in VSTM, then the duration that this representation is maintained in VSTM should be related to the degree to which the representation must be rotated to reach the desired orientation. Therefore, the SPCN should offset at progressively longer latencies as the degree of rotation required increases. We tested this prediction in two experiments utilizing rotated alphanumeric characters. Experiment 1 utilized a normal versus mirror discrimination task that is known to require mental rotation. Experiment 2 utilized a letter versus digit discrimination, a task that does not require mental rotation. In Experiment 1, the offset latency of the SPCN wave increased with increases in the angle of rotation of the target. This effect indicates that targets were maintained in VSTM for longer durations as the angle of rotation increased. Experiment 2 revealed that target orientation did not affect SPCN offset latency when subjects did not adopt a mental rotation strategy, confirming that the effects on the SPCN latency effects observed in Experiment 1 were not due to the mere presentation of rotated patterns. Thus, these two experiments provide clear evidence that mental rotation involves representations maintained in VSTM.

When cross-modal spatial attention fails.

There is now convincing evidence that an involuntary shift of spatial attention to a stimulus in one modality can affect the processing of stimuli in other modalities, but inconsistent findings across different paradigms have led to controversy. Such inconsistencies have important implications for theories of cross-modal attention. The authors investigated why orienting attention to a visual event sometimes influences responses to subsequent sounds and why it sometimes fails to do so. They examined visual-cue-on-auditory-target effects in two paradigms--implicit spatial discrimination (ISD) and orthogonal cuing (OC)--that have yielded conflicting findings in the past. Consistent with previous research, visual cues facilitated responses to same-side auditory targets in the ISD paradigm but not in the OC paradigm. Furthermore, in the ISD paradigm, visual cues facilitated responses to auditory targets only when the targets were presented directly at the cued location, not when they appeared above or below the cued location. This pattern of results confirms recent claims that visual cues fail to influence responses to auditory targets in the OC paradigm because the targets fall outside the focus of attention.

Lateralized readiness potentials reveal motor slowing in the aging brain.

Older adults consistently show slower reaction times (RTs) to the onset of motion. Both cognitive slowing and motor slowing have been suggested as causes of this effect. The lateralized readiness potential (LRP) of the electroencephalogram can be used to separate perceptual and decision processes from motor programming and execution as causes of RT differences. We used the LRP to discern the origin of slowing in RT to motion onset that occurs in elderly individuals. After the onset of motion in a visual display, we asked participants to identify the direction of that motion (up or down) by pressing a button. Older participants showed significantly slower RTs than did younger participants. The LRP showed that the bulk of slowed response arose from slowed motor processes, rather than perceptual processing. We discuss the differences found in amplitude and onset latency of the LRP in the context of theories of motion processing and inhibition in the aging brain.

Cortical expressions of inhibition of return.

Inhibition of return (IOR) is a phenomenon that has been thought to be closely associated with attention mechanisms. In particular, it might arise from the operation of an attentional mechanism that facilitates visual search by inhibiting both covert attention and eye movements from returning to recently inspected locations. Although IOR has received a great deal of research interest, and mechanisms involving sensory, perceptual, and motor consequences have been proposed, no consensus has yet been reached regarding the stages of information processing at which IOR operates. In the present study, we utilized event-related potential (ERP) measures of visual and motor processes to investigate the processing changes underlying IOR. In three experiments, involving localization, detection, or Go-NoGo discrimination, participants were required to make manual responses to target stimuli. In each of these experiments, IOR was associated with a slowing of premotor processes as indicated by a modulation of the onset of the target-locked lateralized readiness potential (LRP). However, the duration of motor processes was not affected (response-locked LRP latency). Consistent with a perceptual locus of IOR, the amplitudes of the occipital ERP peaks were reduced for targets at cued locations relative to those at uncued locations. These and earlier results together provide considerable support for a model in which salience mechanisms that guide attention orienting are also affected by IOR, in that processing a stimulus at a location results in a lowering of its salience for future processing, making orienting to that location, and responding to targets presented there, more time consuming.

Event-related potential evidence for attentional inhibition of return in audition.

Orienting attention to a spatial location facilitates responding to a subsequent target at that location, but inhibits the response if attention is oriented away from that location before the target appears there. This inhibitory effect of attention re-orienting, called inhibition of return (IOR), occurs in vision, hearing, touch, and cross-modally, and has been well studied behaviorally. However, little is known about its underlying neural mechanism(s). We report a study of the neural mechanism of auditory IOR using event-related potentials (ERPs). Auditory IOR was associated with elimination, but not reversal, of the Nd1 difference wave. Previous research indicates that the Nd1 represents an enhanced neural response to attended sounds. Also, auditory IOR was associated with a delay in the latency of the peak of the N1 component of the ERP at parietal sites. These effects are consistent with the accounts if inhibition of attention return that have been proposed for IOR, but are somewhat different from effects found in analogous ERP studies of visual IOR.

Object-substitution masking modulates spatial attention deployment and the encoding of information in visual short-term memory: insights from occipito-parietal ERP components.

If object-substitution masking (OSM) arises from mask representations replacing target representations, OSM should impede the formation of representations in visual short-term memory (VSTM). We utilized event-related potentials to examine the effect of OSM on target processing. An N2pc was observed on trials with delayed-offset masks, indicating that focused attention was directed to the target. The sustained posterior contralateral negativity (SPCN), an index of VSTM storage, was observed in delayed-offset trials only on trials with correct responses. This supports the hypothesis that inaccurate performance on delayed-offset trials arises from a failure to encode the target in VSTM. On co-termination trials, accuracy was high and neither the N2pc nor SPCN was observed. This indicates that, in the absence of masking, the task was accomplished by maintaining a diffuse attentional state that enabled the joint encoding of the potential target items.

Response-selection conflict contributes to inhibition of return.

Here we examined the relationship between inhibition of return (IOR) and response-selection conflict. In two go/no-go and spatial-cueing experiments, we measured the amplitude of the fronto-central N2 event-related potential component to estimate the degree of response-selection conflict for validly cued and invalidly cued targets. When the probability of a go target was high (Experiment 1), both the amplitude of the N2 elicited on no-go trials and the number of false alarm errors were greater on invalid-cue than on valid-cue trials. When the probability of a go target was low (Experiment 2), neither of these effects was observed and the magnitude of the IOR effect was greatly reduced. These results show that a relative response bias toward responding on invalid-cue trials contributes to the IOR reaction time effect when the required response is prepotent.

On the relationship between occipital cortex activity and inhibition of return.

The present study explored the relationship between inhibition of return (IOR) and visual processes by seeking evidence that IOR and changes in event-related potential (ERP) indices of occipital cortex activity covary in response to experimental manipulation. The presence or absence of a central reorienting event was manipulated within the context of a cue-target experiment. When a reorienting event was presented in the interval between cue and target, IOR was accompanied by reductions in the amplitudes of early occipital ERP peaks on validly cued trials relative to invalidly cued trials. When a reorienting event was not presented, neither IOR nor modulations of the occipital ERP peaks was observed. These results provide strong evidence that IOR arises from changes in occipital visual processing. We propose that IOR arises from a slowing of response-selection processes on validly cued trials due differences in the perceptual input to the decision-making process.

Tracking the location of visuospatial attention in a contingent capture paradigm.

Currently, there is considerable controversy regarding the degree to which top-down control can affect attentional capture by salient events. According to the contingent capture hypothesis, attentional capture by a salient stimulus is contingent on a match between the properties of the stimulus and top-down attentional control settings. In contrast, bottom-up saliency accounts argue that the initial capture of attention is determined solely by the relative salience of the stimulus, and the effect of top-down attentional control is limited to effects on the duration of attentional engagement on the capturing stimulus. In the present study, we tested these competing accounts by utilizing the N2pc event-related potential component to track the locus of attention during an attentional capture task. The results were completely consistent with the contingent capture hypothesis: An N2pc wave was elicited only by distractors that possessed the target-defining attribute. In a second experiment, we expanded upon this finding by exploring the effect of target-distractor similarity on the duration that attention dwells at the distractor location. In this experiment, only distractors possessing the target-defining attribute (color) captured visuospatial attention to their location and the N2pc increased in duration and in magnitude when the capture distractor also shared a second target attribute (category membership). Finally, in three additional control experiments, we replicated the finding of an N2pc generated by distractors, only if they shared the target-defining attribute. Thus, our results demonstrate that attentional control settings influence both which stimuli attract attention and to what extent they are processed.

Reorienting attention and inhibition of return.

In experiments examining inhibition of return (IOR), it is common practice to present a second cue at fixation during the cue-target interval. The purpose of this fixation cue is to reorient attention away from the cued location to ensure that the facilitative effects of spatial attention do not obscure IOR. However, despite their frequent use, relatively little is known about the relationship between fixation cues and IOR. In the present experiments, we examined the role of fixation cues by manipulating their presence in tasks that either did or did not require target identification. When the participants were required to either detect (Experiment 1A) or localize (Experiment 2A) a target, the magnitude of IOR was unaffected by the presence of a fixation cue. In contrast, when the participants were required to identify a target (Experiments 1B, 2B, and 3), IOR was observed only when a fixation cue was presented. This result was obtained regardless of the type of response that was required (two-alternative forced choice or go/no go). The effectiveness of the fixation cue in revealing IOR in these tasks is consistent with its putative role in reorienting attention away from the cued location.

Inhibition of return and response repetition within and between modalities.

Inhibition of return (IOR) refers to slower responding to stimuli at previously occupied spatial locations. IOR has been vigorously studied because of its possible deep involvement with attention mechanisms. Although IOR occurs both within and across modalities in several experimental paradigms for simple stimulus detection tasks, it has sometimes been difficult to demonstrate in perceptual discrimination tasks. In the preferred target-target paradigm, in which responses are made to a series of targets that vary in spatial location, failure to find IOR could possibly result from mixing of spatial IOR with the facilitating effects of stimulus and/or response repetition on discrimination response times. In this paper we report the first demonstration of auditory/auditory and cross-modality IOR in a target-target paradigm using a discrimination task. Our results show that IOR occurs in this task only on trials on which stimuli and responses are not repeated. These findings present a challenge to purely visual accounts of IOR and support the view that IOR arises within a more general, supra-modal mechanism of attention.

Inhibition of return from stimulus to response.

In a standard inhibition-of-return (IOR) paradigm using a manual key-press response, we examined the effect of IOR both on the amplitude of early sensory event-related brain potential (ERP) components and on the motor-related lateralized readiness potential (LRP). IOR was associated with a delay of premotor processes (target-locked LRP latency) and reduced sensory ERP activity. No effect of IOR was found on motor processes (response-locked LRP latency). Thus, IOR must arise at least in part from changes in perceptual processes, and, at least when measured with manual key presses, IOR does not arise from inhibition of motor processes. These results are consistent with the results of attention-orienting studies and provide support for an inhibition-of-attention explanation for IOR.

Auditory frequency-based inhibition differs from spatial IOR.

Uninformative auditory frequency cues have a facilitatory effect on reaction time and accuracy of detection and intensity discrimination of target tones for cue-target intervals of up to 3 sec (Green & McKeown, 2001; Ward, 1997). Under some conditions, however, this facilitatory effect can reverse to an inhibitory effect at cue-target intervals longer than 450 msec (Mondor, Breau, & Milliken, 1998). Thepresent work demonstrates that such inhibitory effects are not found in target-target experiments (Experiment 1) or in cue-target experiments requiring a go-no-go discrimination of the target (Experiment 2), whereas they do appear in the paradigm used by Mondor et al. (1998, Experiment 3), albeit unaffected by the similarity of cue and target. Thus, the frequency-based inhibitory effects sometimes found in auditory cuing tasks can be distinguished empirically from those characterizing spatial inhibition of return (IOR), which are found in both target-target and go-no-go cue-target paradigms. The present work and functional and neurophysiological arguments all support the position that different mechanisms underlie spatial IOR and the inhibitory effects sometimes found in auditory frequency processing.