John J. McDonald

Electrophysiological Evidence of the Capture of Visual Attention

We investigated the ability of salient yet task-irrelevant stimuli to capture attention in two visual search experiments. Participants were presented with circular search arrays that contained a highly salient distractor singleton defined by color and a less salient target singleton defined by form. A component of the event-related potential called the N2pc was used to track the allocation of attention to lateralized positions in the arrays. In Experiment 1, a lateralized distractor elicited an N2pc when a concurrent target was presented on the vertical meridian and thus could not elicit lateralized components such as the N2pc. A similar distractor-elicited N2pc was found in Experiment 2, which was conducted to rule out certain voluntary search strategies. Additionally, in Experiment 2 both the distractor and the target elicited the N2pc component when the two stimuli were presented on opposite sides of the search array. Critically, the distractor-elicited N2pc preceded the target-elicited N2pc on these trials. These results demonstrate that participants shifted attention to the target only after shifting attention to the more salient but task-irrelevant distractor. This pattern of results is in line with theories of attention in which stimulus-driven control plays an integral role.

Electrophysiological indices of target and distractor processing in visual search

Attentional selection of a target presented among distractors can be indexed with an event-related potential (ERP) component known as the N2pc. Theoretical interpretation of the N2pc has suggested that it reflects a fundamental mechanism of attention that shelters the cortical representation of targets by suppressing neural activity stemming from distractors. Results from fields other than human electrophysiology, however, suggest that attention does not act solely through distractor suppression; rather, it modulates the processing of both target and distractors. We conducted four ERP experiments designed to investigate whether the N2pc reflects multiple attentional mechanisms. Our goal was to reconcile ostensibly conflicting outcomes obtained in electrophysiological studies of attention with those obtained using other methodologies. Participants viewed visual search arrays containing one target and one distractor. In Experiments 1 through 3, the distractor was isoluminant with the background, and therefore, did not elicit early lateralized ERP activity. This work revealed a novel contralateral ERP component that appears to reflect direct suppression of the cortical representation of the distractor. We accordingly name this component the distractor positivity (PD). In Experiment 4, an ERP component associated with target processing was additionally isolated. We refer to this component as the target negativity (NT). We believe that the N2pc reflects the summation of the PD and NT, and that these discrete components may have been confounded in earlier electrophysiological studies. Overall, this study demonstrates that attention acts on both target and distractor representations, and that this can be indexed in the visual ERP.

Performance monitoring in the anterior cingulate is not all error related: Expectancy deviation and the representation of action-outcome associations

Several converging lines of evidence suggest that the anterior cingulate cortex (ACC) is selectively involved in error detection or evaluation of poor performance. Here we challenge this notion by presenting event-related potential (ERP) evidence that the feedback-elicited error-related negativity, an ERP component attributed to the ACC, can be elicited by positive feedback when a person is expecting negative feedback and vice versa. These results suggest that performance monitoring in the ACC is not limited to error processing. We propose that the ACC acts as part of a more general performance-monitoring system that is activated by violations in expectancy. Further, we propose that the common observation of increased ACC activity elicited by negative events could be explained by an overoptimistic bias in generating expectations of performance. These results could shed light into neurobehavioral disorders, such as depression and mania, associated with alterations in performance monitoring and also in judgments of self-related events.

Effects of Spatial Congruity on Audio-Visual Multimodal Integration

Spatial constraints on multisensory integration of auditory (A) and visual (V) stimuli were investigated in humans using behavioral and electrophysiological measures. The aim was to find out whether cross-modal interactions between A and V stimuli depend on their spatial congruity, as has been found for multisensory neurons in animal studies (Stein & Meredith, 1993). Randomized sequences of unimodal (A or V) and simultaneous bimodal (AV) stimuli were presented to right-or left-field locations while subjects made speeded responses to infrequent targets of greater intensity that occurred in either or both modalities. Behavioral responses to the bimodal stimuli were faster and more accurate than to the uni-modal stimuli for both same-location and different-location AV pairings. The neural basis of this cross-modal facilitation was studied by comparing event-related potentials (ERPs) to the bimodal AV stimuli with the summed ERPs to the unimodal A and V stimuli. These comparisons revealed neural interactions localized to the ventral occipito-temporal cortex (at 190 msec) and to the superior temporal cortical areas (at 260 msec) for both same-and different-location AV pairings. In contrast, ERP interactions that differed according to spatial congruity included a phase and amplitude modulation of visual-evoked activity localized to the ventral occipito-temporal cortex at 100-400 msec and an amplitude modulation of activity localized to the superior temporal region at 260-280 msec. These results demonstrate overlapping but distinctive patterns of multisensory integration for spatially congruent and incongruent AV stimuli.

Neural Substrates of Perceptual Enhancement by Cross-Modal Spatial Attention

Orienting attention involuntarily to the location of a sudden sound improves perception of subsequent visual stimuli that appear nearby. The neural substrates of this cross-modal attention effect were investigated by recording event-related potentials to the visual stimuli using a dense electrode array and localizing their brain sources through inverse dipole modeling. A spatially nonpredictive auditory precue modulated visual-evoked neural activity first in the superior temporal cortex at 120140 msec and then in the ventral occipital cortex of the fusiform gyrus 1525 msec later. This spatio-temporal sequence of brain activity suggests that enhanced visual perception produced by the cross-modal orienting of spatial attention results from neural feedback from the multimodal superior temporal cortex to the visual cortex of the ventral processing stream.

Involuntary Listening Aids Seeing: Evidence From Human Electrophysiology

It is well known that sensory events of one modality can influence judgments of sensory events in other modalities. For example, people respond more quickly to a target appearing at the location of a previous cue than to a target appearing at another location, even when the two stimuli are from different modalities. Such cross-modal interactions suggest that involuntary spatial attention mechanisms are not entirely modality-specific. In the present study, event-related brain potentials (ERPs) were recorded to elucidate the neural basis and timing of involuntary, cross-modal spatial attention effects. We found that orienting spatial attention to an irrelevant sound modulates the ERP to a subsequent visual target over modality-specific, extrastriate visual cortex, but only after the initial stages of sensory processing are completed. These findings are consistent with the proposal that involuntary spatial attention orienting to auditory and visual stimuli involves shared, or at least linked, brain mechanisms.

Rhythms of Consciousness: Binocular Rivalry Reveals Large-Scale Oscillatory Network Dynamics Mediating Visual Perception

Consciousness has been proposed to emerge from functionally integrated large-scale ensembles of gamma-synchronous neural populations that form and dissolve at a frequency in the theta band. We propose that discrete moments of perceptual experience are implemented by transient gamma-band synchronization of relevant cortical regions, and that disintegration and reintegration of these assemblies is time-locked to ongoing theta oscillations. In support of this hypothesis we provide evidence that (1) perceptual switching during binocular rivalry is time-locked to gamma-band synchronizations which recur at a theta rate, indicating that the onset of new conscious percepts coincides with the emergence of a new gamma-synchronous assembly that is locked to an ongoing theta rhythm; (2) localization of the generators of these gamma rhythms reveals recurrent prefrontal and parietal sources; (3) theta modulation of gamma-band synchronization is observed between and within the activated brain regions. These results suggest that ongoing theta-modulated-gamma mechanisms periodically reintegrate a large-scale prefrontal-parietal network critical for perceptual experience. Moreover, activation and network inclusion of inferior temporal cortex and motor cortex uniquely occurs on the cycle immediately preceding responses signaling perceptual switching. This suggests that the essential prefrontal-parietal oscillatory network is expanded to include additional cortical regions relevant to tasks and perceptions furnishing consciousness at that moment, in this case image processing and response initiation, and that these activations occur within a time frame consistent with the notion that conscious processes directly affect behaviour.

Neural basis of auditory-induced shifts in visual time-order perception

Attended objects are perceived to occur before unattended objects even when the two objects are presented simultaneously. This finding has led to the widespread view that attention modulates the speed of neural transmission in the various perceptual pathways. We recorded event-related potentials during a time-order judgment task to determine whether a reflexive shift of attention to a sudden sound modulates the speed of sensory processing in the human visual system. Attentional cueing influenced the perceived order of lateralized visual events but not the timing of event-related potentials in visual cortex. Attentional cueing did, however, enhance the amplitude of neural activity in visual cortex, which shows that attention-induced shifts in visual time-order perception can arise from modulations of signal strength rather than processing speed in the early visual-cortical pathways.

Cross-modal cueing of attention alters appearance and early cortical processing of visual stimuli

The question of whether attention makes sensory impressions appear more intense has been a matter of debate for over a century. Recent psychophysical studies have reported that attention increases apparent contrast of visual stimuli, but the issue continues to be debated. We obtained converging neurophysiological evidence from human observers as they judged the relative contrast of visual stimuli presented to the left and right visual fields following a lateralized auditory cue. Cross-modal cueing of attention boosted the apparent contrast of the visual target in association with an enlarged neural response in the contralateral visual cortex that began within 100 ms after target onset. The magnitude of the enhanced neural response was positively correlated with perceptual reports of the cued target being higher in contrast. The results suggest that attention increases the perceived contrast of visual stimuli by boosting early sensory processing in the visual cortex.

Electrical Neuroimaging Reveals Timing of Attentional Control Activity in Human Brain

Voluntarily shifting attention to a location of the visual field improves the perception of events that occur there. Regions of frontal cortex are thought to provide the top-down control signal that initiates a shift of attention, but because of the temporal limitations of functional brain imaging, the timing and sequence of attentional-control operations remain unknown. We used a new analytical technique (beamformer spatial filtering) to reconstruct the anatomical sources of low-frequency brain waves in humans associated with attentional control across time. Following a signal to shift attention, control activity was seen in parietal cortex 100–200 ms before activity was seen in frontal cortex. Parietal cortex was then reactivated prior to anticipatory biasing of activity in occipital cortex. The magnitudes of early parietal activations were strongly predictive of the degree of attentional improvement in perceptual performance. These results show that parietal cortex, not frontal cortex, provides the initial signals to shift attention and indicate that top-down attentional control is not purely top down.