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S A Hillyard

Publications and source records attributed to S A Hillyard.

At least 37 records · Page 2Linked to original sources

Sensory gain control (amplification) as a mechanism of selective attention: electrophysiological and neuroimaging evidence.

Both physiological and behavioral studies have suggested that stimulus-driven neural activity in the sensory pathways can be modulated in amplitude during selective attention. Recordings of event-related brain potentials indicate that such sensory gain control or amplification processes play an important role in visual-spatial attention. Combined event-related brain potential and neuroimaging experiments provide strong evidence that attentional gain control operates at an early stage of visual processing in extrastriate cortical areas. These data support early selection theories of attention and provide a basis for distinguishing between separate mechanisms of attentional suppression (of unattended inputs) and attentional facilitation (of attended inputs).

Attention↗

Event-related brain potentials in the study of visual selective attention.

Event-related brain potentials (ERPs) provide high-resolution measures of the time course of neuronal activity patterns associated with perceptual and cognitive processes. New techniques for ERP source analysis and comparisons with data from blood-flow neuroimaging studies enable improved localization of cortical activity during visual selective attention. ERP modulations during spatial attention point toward a mechanism of gain control over information flow in extrastriate visual cortical pathways, starting about 80 ms after stimulus onset. Paying attention to nonspatial features such as color, motion, or shape is manifested by qualitatively different ERP patterns in multiple cortical areas that begin with latencies of 100-150 ms. The processing of nonspatial features seems to be contingent upon the prior selection of location, consistent with early selection theories of attention and with the hypothesis that spatial attention is "special."

Animals↗

Spatio-temporal dynamics of attention to color: evidence from human electrophysiology.

This study characterized patterns of brain electrical activity associated with selective attention to the color of a stimulus. Multichannel recordings of event-related potentials (ERPs) were obtained while subjects viewed randomized sequences of checkerboards consisting of isoluminant red or blue checks superimposed on a grey background. Stimuli were presented foveally at a rapid rate, and subjects were required to attend to the red or blue checks in separate blocks of trials and to press a button each time they detected a dimmer target stimulus of the attended color. An early negative ERP component with an onset latency of 50 ms was sensitive to stimulus color but was unaffected by the attentional manipulation. ERPs elicited by attended and unattended stimuli began to diverge after approximately 100 ms following stimulus onset. Inverse dipole modelling of the attended-minus-unattended difference waveform indicated that an initial positive deflection with an onset latency of 100 ms had a source in lateral occipital cortex, while a subsequent negative deflection with an onset at 160 ms had a source in inferior occipito-temporal cortex. Longer-latency attention-sensitive components were localized to premotor frontal areas (onset at 190 ms) and to more anterior regions of the fusiform gyrus (onset at 240 ms). These source localizations correspond closely with cortical areas that were identified in previous neuroimaging studies as being involved in color-selective processing. The present ERP data thus provide information about the time course of stimulus selection processes in cortical areas that subserve attention to color.

Adult↗

Effects of spatial selective attention on the steady-state visual evoked potential in the 20-28 Hz range.

Steady-state visual evoked potentials (SSVEPs) were recorded from the scalp of subjects who attended to a flickering LED display in one visual field while ignoring a similar display (flickering at a different frequency) in the opposite visual field. The flicker frequencies were 20.8 Hz in the left-field display and 27.8 Hz in the right-field display. The SSVEP to the flicker in either field was enhanced in amplitude when attention was directed to its location. The scalp distribution of this SSVEP enhancement was narrowly focused over the posterior scalp contralateral to the visual field of stimulation. A source analysis using Variable Resolution Electromagnetic Tomography (VARETA) indicated that the source current densities for the SSVEP attention effect had a focal origin in the contralateral parieto-occipital cortex.

Adolescent↗

Temporal dynamics of early perceptual processing.

Recordings of electrical and magnetic brain responses to sensory stimulation provide high-resolution measures of the time course of early perceptual processing. Spatio-temporal analyses of brain activity patterns during the first 200 ms after stimulus presentation have characterized the timing of attentional selection processes and different stages of feature encoding and pattern analyses. Recent studies that incorporate blood flow neuroimaging techniques provide support for mechanisms of early selection of attended visual inputs in extrastriate cortical pathways. The spatial tuning properties of early auditory selection have also been delineated. Electrical and magnetic responses that index the encoding of higher-order pattern information have been identified in both visual and auditory modalities and localized to specific cortical areas.

Attention↗

Magnetoencephalographic recordings demonstrate attentional modulation of mismatch-related neural activity in human auditory cortex.

It is widely agreed that the negative brain potential elicited at 150-200 ms by a deviant, less intense sound in a repetitive series can be modulated by attention. To investigate whether this modulation represents a genuine attention effect on the mismatch negativity (MMN) arising from auditory cortex or attention-related activity from another brain region, we recorded both the MMN and the mismatch magnetic field (MMF) elicited by such deviants in a dichotic listening task. Deviant tones in the attended ear elicited a sizable MMF that was well modeled as a dipolar source in auditory cortex. Both the MMN and MMF to unattended-ear deviants were highly attenuated. These findings support the view that the MMN/MMF elicited in auditory cortex by intensity deviants, and thus the underlying feature-analysis and mismatch-detection processes, are not strongly automatic but rather can be gated or suppressed if attention is strongly focused elsewhere.

Acoustic Stimulation↗

The time course of cortical facilitation during cued shifts of spatial attention.

Adaptive behavior requires the rapid switching of attention among potentially relevant stimuli that appear in the environment. The present study used an electrophysiological approach to continuously measure the time course of visual pathway facilitation in human subjects as attention was shifted from one location to another. Steady-state visual evoked potentials (SSVEPs) were recorded to rapidly flickering lights at attended and unattended locations, and variations in SSVEP amplitude over time were calculated after a cue to shift attention. The build-up of cortical facilitation reflected in SSVEP amplitude was found to bear a close temporal relationship with the emergence of accurate target discriminations at the newly attended location.

Attention↗

The gradient of spatial auditory attention in free field: an event-related potential study.

Young adult subjects attended selectively to brief noise bursts delivered in free field via a horizontal array of seven loudspeakers spaced apart by 9 degrees of angle. Frequent "standard" stimuli (90%) and infrequent "target/deviant" stimuli (10%) of increased bandwidth were delivered at a fast rate in a random sequence equiprobably from each speaker. In separate runs, the subjects' task was to selectively attend to the leftmost, center, or rightmost speaker and to press a button to the infrequent "target" stimuli occurring at the designated spatial location. Behavioral detection rates and concurrently recorded event-related potentials (ERPs) indicated that auditory attention was deployed as a finely tuned gradient around the attended sound source, thus providing support for gradient models of auditory spatial attention. Furthermore, the ERP data suggested that the spatial focusing of attention was achieved in two distinct stages, with an early more broadly tuned filtering of inputs occurring over the first 80-200 msec after stimulus onset, followed by a more narrowly focused selection of attended-location deviants that began at around 250 msec and closely resembled the behavioral gradient of target detections.

Acoustic Stimulation↗

Magnetoencephalographic recording of steady-state visual evoked cortical activity.

Steady-state visual evoked magnetic fields (SSVEFs) were recorded in response to a flickering light source using a 37-channel magnetometer. The SSVEF had a sinusoidal waveform having the same fundamental frequency as the driving stimulus, which was either 6.0 Hz, 11.9 Hz, or 15.2 Hz. SSVEF topographies at each frequency had a dipoloar form over the posterior head that could well-modelled by single equivalent current dipoles. The best-fit dipoles were localized in posterior occipital cortex for the SSVEFs to 6.0 and 11.9 Hz stimuli and in more anterior and ventromedial occipital cortex for the 15.2 Hz SSVEP.

Adult↗

Neural mechanisms of spatial selective attention in areas V1, V2, and V4 of macaque visual cortex.

Many neurons in extrastriate visual cortex have large receptive fields, and this may lead to significant computational problems whenever multiple stimuli fall within a single field. Previous studies have suggested that when multiple stimuli fall within a cell's receptive field, they compete for the cell's response in a manner that can be biased in favor of attended stimuli. In the present study we examined this role of attention in areas V1, V2, and V4 of macaque monkeys with the use of a behavioral paradigm in which attention was directed to one of two stimulus locations. When two stimuli were presented simultaneously inside the cell's receptive field (which could be accomplished only in areas V2 and V4), we found that the cell's response was strongly influenced by which of the two stimuli was attended. The size of this attention effect was reduced when the attended and ignored stimuli were presented sequentially rather than simultaneously. In addition, the effects became very weak and inconsistent in these areas when only one of the two stimuli was located inside the receptive field. Attention thus modulated sensory responses primarily when two or more simultaneous stimuli competed for access to a neuron's receptive field. As in areas V2 and V4, attention did not modulate sensory responses in area V1 when only a single stimulus was inside the receptive field. In addition, the small receptive fields in this area precluded the simultaneous presentation of attended and ignored stimuli inside the receptive field, making it impossible to determine whether attention effects would be observed under the conditions that led to consistent attention effects in areas V2 and V4. Spontaneous firing rates in areas V2 and V4 were found to be 30-40% higher when attention was directed inside rather than outside the receptive field, even when no stimulus was present in the receptive field. Spontaneous firing rates also varied according to the particular location within the receptive field that was attended. These shifts in spontaneous activity may reflect a top-down signal that biases responses in favor of stimuli at the attended location.

Animals↗

Selective attention to stimulus location modulates the steady-state visual evoked potential.

Steady-state visual evoked potentials (SSVEPs) were recorded from the scalp of human subjects who were cued to attend to a rapid sequence of alphanumeric characters presented to one visual half-field while ignoring a concurrent sequence of characters in the opposite half-field. These two-character sequences were each superimposed upon a small square background that was flickered at a rate of 8.6 Hz in one half-field and 12 Hz in the other half-field. The amplitude of the frequency-coded SSVEP elicited by either of the task-irrelevant flickering backgrounds was significantly enlarged when attention was focused upon the character sequence at the same location. This amplitude enhancement with attention was most prominent over occipital-temporal scalp areas of the right cerebral hemisphere regardless of the visual field of stimulation. These findings indicate that the SSVEP reflects an enhancement of neural responses to all stimuli that fall within the "spotlight" of spatial attention, whether or not the stimuli are task-relevant. Recordings of the SSVEP provide a new approach for studying the neural mechanisms and functional properties of selective attention to multi-element visual displays.

Adult↗

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.

Attention↗

Selective attention to the color and direction of moving stimuli: electrophysiological correlates of hierarchical feature selection.

Event-related brain potentials (ERPs) were recorded from subjects who attended to pairs of adjacent colored squares that were flashed sequentially to produce a perception of movement. The task was to attend selectively to stimuli in one visual field and to detect slower moving targets that contained the critical value of the attended feature, be it color or movement direction. Attention to location was reflected by a modulation of the early P1 and N1 components of the ERP, whereas selection of the relevant stimulus feature was associated with later selection negativity components. ERP indices of feature selection were elicited only by stimuli at the attended location and had distinctive scalp distributions for features mediated by "ventral" (color) and "dorsal" (motion) cortical areas. ERP indices of target selection were also contingent on the prior selection of location but initially did not depend on the selection of the relevant feature. These ERP data reveal the timing of sequential, parallel, and contingent stages of visual processing and support early-selection theories of attention that stipulate attentional control over the initial processing of stimulus features.

Adolescent↗

Semantic processing and memory for attended and unattended words in dichotic listening: behavioral and electrophysiological evidence.

Thirty-two Ss studied words presented to 1 ear, while ignoring a concurrent word list presented to the opposite ear. The N400 component of the event-related potentials elicited by attended words was modulated by semantic priming between successive words. The N400 elicited by unattended words was insensitive to semantic manipulation. Recognition memory was better for attended than for unattended words. However, the percentage of false positives was elevated equally for lures that were semantically related to "old" words, whether they had been attended or unattended. Words that were initially attended induced similar repetition effects in a lexical decision task as words that were initially unattended. Hence, both attended and unattended words are semantically processed and activate semantic representations. However, attended words from traces that are subsequently more available to conscious recollection than unattended words.

Adult↗

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.

Attention↗

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.

Brain↗

Sources of attention-sensitive visual event-related potentials.

In a study of the neural processes that mediate visual attention in humans, 32-channel recordings of event-related potentials were obtained from 14 normal subjects while they performed a spatial attention task. The generator locations of the early C1, P1, and N1 components of the visual evoked response were estimated by means of topographic maps of voltage and current source density in conjunction with dipole modelling. The topography of the C1 component (ca. 85 ms post-stimulus) was consistent with a generator in striate cortex, and this component was unaffected by attention. In contrast, the P1 and N1 components (ca. 95 and 170 ms) exhibited current density foci at scalp sites overlying lateral extrastriate cortex and were larger for attended stimuli than for unattended stimuli. The voltage topographies in the 75-175 ms latency range were modeled with a 5-dipole configuration consisting of a single striate dipole and left-right pairs of dipoles located in lateral extrastriate and inferior occipito-temporal areas. This model was found to account for the voltage topographies produced by both attended and unattended stimuli with low residual variance. These results support the proposal that visual-spatial attention modulates neural activity in extrastriate visual cortex but does not affect the initial evoked response in striate cortex.

Adolescent↗

Effects of aging on event-related brain potentials (ERPs) in a visual detection task.

Event-related brain potentials (ERPs) were recorded from 74 subjects (45 men) between 18 and 82 years of age in a simple visual detection task. On each trial the subject reported the location of a triangular flash of light presented briefly 20 degrees laterally to the left or right visual field or to both fields simultaneously. ERPs to targets exhibited a similar morphology including P1, N1, P2, N2, and P3 components across all age groups. The principal effects of advancing age were (1) a marked reduction in amplitude of the posterior P1 component (75-150 latency) together with an amplitude increase of an anterior positivity at the same latency; (2) an increase in amplitude of the P3 component that was most prominent over frontal scalp areas; and (3) a linear increase in P3 peak latency. These results extend the findings of age-related changes in P3 peak latency and distribution to a non-oddball task in the visual modality and raise the possibility that short-latency ERPs may index changes in visual attention in the elderly.

Adolescent↗