The time course of feature-selective attention inside and outside the focus of spatial attention.

Research on attentional selection of stimulus features has yielded seemingly contradictory results. On the one hand, many experiments in humans and animals have observed a "global" facilitation of attended features across the entire visual field, even when spatial attention is focused on a single location. On the other hand, several event-related potential studies in humans reported that attended features are enhanced at the attended location only. The present experiment demonstrates that these conflicting results can be explained by differences in the timing of attentional allocation inside and outside the spatial focus of attention. Participants attended to fields of either red or blue randomly moving dots on either the left or right side of fixation with the task of detecting brief coherent motion targets. Recordings of steady-state visual evoked potentials elicited by the flickering stimuli allowed concurrent measurement of the time course of feature-selective attention in visual cortex on both the attended and the unattended sides. The onset of feature-selective attentional modulation on the attended side occurred around 150 ms earlier than on the unattended side. This finding that feature-selective attention is not spatially global from the outset but extends to unattended locations after a temporal delay resolves previous contradictions between studies finding global versus hierarchical selection of features and provides insight into the fundamental relationship between feature-based and location-based (spatial) attention mechanisms.

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.

Split-Brain: What We Know Now and Why This is Important for Understanding Consciousness.

Recently, the discussion regarding the consequences of cutting the corpus callosum ("split-brain") has regained momentum (Corballis, Corballis, Berlucchi, & Marzi, Brain, 141(6), e46, 2018; Pinto et al., Brain, 140(5), 1231-1237, 2017a; Pinto, Lamme, & de Haan, Brain, 140(11), e68, 2017; Volz & Gazzaniga, Brain, 140(7), 2051-2060, 2017; Volz, Hillyard, Miller, & Gazzaniga, Brain, 141(3), e15, 2018). This collective review paper aims to summarize the empirical common ground, to delineate the different interpretations, and to identify the remaining questions. In short, callosotomy leads to a broad breakdown of functional integration ranging from perception to attention. However, the breakdown is not absolute as several processes, such as action control, seem to remain unified. Disagreement exists about the responsible mechanisms for this remaining unity. The main issue concerns the first-person perspective of a split-brain patient. Does a split-brain harbor a split consciousness or is consciousness unified? The current consensus is that the body of evidence is insufficient to answer this question, and different suggestions are made with respect to how future studies might address this paucity. In addition, it is suggested that the answers might not be a simple yes or no but that intermediate conceptualizations need to be considered.

Neural Correlates of Enhanced Visual Attentional Control in Action Video Game Players: An Event-Related Potential Study.

Action video game players (AVGPs) outperform non-action video game players (NAVGPs) on a range of perceptual and attentional tasks. Although several studies have reported neuroplastic changes within the frontoparietal networks of attention in AVGPs, little is known about possible changes in attentional modulation in low-level visual areas. To assess the contribution of these different levels of neural processing to the perceptual and attentional enhancements noted in AVGPs, visual event-related potentials (ERPs) were recorded from 14 AVGPs and 14 NAVGPs during a target discrimination task that required participants to attend to rapid sequences of Gabor patches under either focused or divided attention conditions. AVGPs responded faster to target Gabors in the focused attention condition compared with the NAVGPs. Correspondingly, ERPs to standard Gabors revealed a more pronounced negativity in the time range of the parietally generated anterior N1 component in AVGPs compared with NAVGPs during focused attention. In addition, the P2 component of the visual ERP was more pronounced in AVGPs than in NAVGPs over the hemisphere contralateral to the stimulus position in response to standard Gabors. Contrary to predictions, however, attention-modulated occipital components generated in the low-level extrastriate visual pathways, including the P1 and posterior N1, showed no significant group differences. Thus, the main neural signature of enhanced perceptual and attentional control functions in AVGPs appears linked to an attention-dependent parietal process, indexed by the anterior N1 component, and possibly to more efficient higher-order perceptual processing, indexed by the P2 component.

Neural mechanisms of divided feature-selective attention to colour.

Feature-based attentional selection of colour is challenging to investigate due to the multidimensional nature of colour-space. When attending concurrently to features from different feature dimensions (e.g. red and horizontal), the attentional selections of the separate dimensions are largely independent. Therefore, if colour constitutes multiple independent feature dimensions for attentional purposes, concurrently attending to two colours should be effective and independent of the specific configuration of target and distractor colours. Here, observers attended concurrently to two out of four fully overlapping random dot kinematograms of different colours, and the allocation of attention to each colour was assessed separately by recordings of steady-state visual evoked potentials. The magnitude of attention effects depended on colour proximity and was well described by a simple model which suggested that colour space is rescaled in an adaptive manner to achieve attentional selection. In conclusion, different spatially overlaid colours can be attended concurrently with an efficiency that is determined by their configuration in colour space, supporting the idea that (at least in terms of hue) colour acts as a single dimension for attentional purposes.

Perceptual Learning Induces Persistent Attentional Capture by Nonsalient Shapes.

Visual attention can be attracted automatically by salient simple features, but whether and how nonsalient complex stimuli such as shapes may capture attention in humans remains unclear. Here, we present strong electrophysiological evidence that a nonsalient shape presented among similar shapes can provoke a robust and persistent capture of attention as a consequence of extensive training in visual search (VS) for that shape. Strikingly, this attentional capture that followed perceptual learning (PL) was evident even when the trained shape was task-irrelevant, was presented outside the focus of top-down spatial attention, and was undetected by the observer. Moreover, this attentional capture persisted for at least 3-5 months after training had been terminated. This involuntary capture of attention was indexed by electrophysiological recordings of the N2pc component of the event-related brain potential, which was localized to ventral extrastriate visual cortex, and was highly predictive of stimulus-specific improvement in VS ability following PL. These findings provide the first evidence that nonsalient shapes can capture visual attention automatically following PL and challenge the prominent view that detection of feature conjunctions requires top-down focal attention.

Involuntary orienting of attention to a sound desynchronizes the occipital alpha rhythm and improves visual perception.

Directing attention voluntarily to the location of a visual target results in an amplitude reduction (desynchronization) of the occipital alpha rhythm (8-14Hz), which is predictive of improved perceptual processing of the target. Here we investigated whether modulations of the occipital alpha rhythm triggered by the involuntary orienting of attention to a salient but spatially non-predictive sound would similarly influence perception of a subsequent visual target. Target discrimination was more accurate when a sound preceded the target at the same location (validly cued trials) than when the sound was on the side opposite to the target (invalidly cued trials). This behavioral effect was accompanied by a sound-induced desynchronization of the alpha rhythm over the lateral occipital scalp. The magnitude of alpha desynchronization over the hemisphere contralateral to the sound predicted correct discriminations of validly cued targets but not of invalidly cued targets. These results support the conclusion that cue-induced alpha desynchronization over the occipital cortex is a manifestation of a general priming mechanism that improves visual processing and that this mechanism can be activated either by the voluntary or involuntary orienting of attention. Further, the observed pattern of alpha modulations preceding correct and incorrect discriminations of valid and invalid targets suggests that involuntary orienting to the non-predictive sound has a rapid and purely facilitatory influence on processing targets on the cued side, with no inhibitory influence on targets on the opposite side.

Attentional Selection of Feature Conjunctions Is Accomplished by Parallel and Independent Selection of Single Features.

Experiments that study feature-based attention have often examined situations in which selection is based on a single feature (e.g., the color red). However, in more complex situations relevant stimuli may not be set apart from other stimuli by a single defining property but by a specific combination of features. Here, we examined sustained attentional selection of stimuli defined by conjunctions of color and orientation. Human observers attended to one out of four concurrently presented superimposed fields of randomly moving horizontal or vertical bars of red or blue color to detect brief intervals of coherent motion. Selective stimulus processing in early visual cortex was assessed by recordings of steady-state visual evoked potentials (SSVEPs) elicited by each of the flickering fields of stimuli. We directly contrasted attentional selection of single features and feature conjunctions and found that SSVEP amplitudes on conditions in which selection was based on a single feature only (color or orientation) exactly predicted the magnitude of attentional enhancement of SSVEPs when attending to a conjunction of both features. Furthermore, enhanced SSVEP amplitudes elicited by attended stimuli were accompanied by equivalent reductions of SSVEP amplitudes elicited by unattended stimuli in all cases. We conclude that attentional selection of a feature-conjunction stimulus is accomplished by the parallel and independent facilitation of its constituent feature dimensions in early visual cortex. SIGNIFICANCE STATEMENT: The ability to perceive the world is limited by the brain's processing capacity. Attention affords adaptive behavior by selectively prioritizing processing of relevant stimuli based on their features (location, color, orientation, etc.). We found that attentional mechanisms for selection of different features belonging to the same object operate independently and in parallel: concurrent attentional selection of two stimulus features is simply the sum of attending to each of those features separately. This result is key to understanding attentional selection in complex (natural) scenes, where relevant stimuli are likely to be defined by a combination of stimulus features.

Sounds activate visual cortex and improve visual discrimination.

A recent study in humans (McDonald et al., 2013) found that peripheral, task-irrelevant sounds activated contralateral visual cortex automatically as revealed by an auditory-evoked contralateral occipital positivity (ACOP) recorded from the scalp. The present study investigated the functional significance of this cross-modal activation of visual cortex, in particular whether the sound-evoked ACOP is predictive of improved perceptual processing of a subsequent visual target. A trial-by-trial analysis showed that the ACOP amplitude was markedly larger preceding correct than incorrect pattern discriminations of visual targets that were colocalized with the preceding sound. Dipole modeling of the scalp topography of the ACOP localized its neural generators to the ventrolateral extrastriate visual cortex. These results provide direct evidence that the cross-modal activation of contralateral visual cortex by a spatially nonpredictive but salient sound facilitates the discriminative processing of a subsequent visual target event at the location of the sound. Recordings of event-related potentials to the targets support the hypothesis that the ACOP is a neural consequence of the automatic orienting of visual attention to the location of the sound.

Peripheral sounds rapidly activate visual cortex: evidence from electrocorticography.

Neurophysiological studies with animals suggest that sounds modulate activity in primary visual cortex in the presence of concurrent visual stimulation. Noninvasive neuroimaging studies in humans have similarly shown that sounds modulate activity in visual areas even in the absence of visual stimuli or visual task demands. However, the spatial and temporal limitations of these noninvasive methods prevent the determination of how rapidly sounds activate early visual cortex and what information about the sounds is relayed there. Using spatially and temporally precise measures of local synaptic activity acquired from depth electrodes in humans, we demonstrate that peripherally presented sounds evoke activity in the anterior portion of the contralateral, but not ipsilateral, calcarine sulcus within 28 ms of sound onset. These results suggest that auditory stimuli rapidly evoke spatially specific activity in visual cortex even in the absence of concurrent visual stimulation or visual task demands. This rapid auditory-evoked activation of primary visual cortex is likely to be mediated by subcortical pathways or direct cortical projections from auditory to visual areas.

Global facilitation of attended features is obligatory and restricts divided attention.

In many common situations such as driving an automobile it is advantageous to attend concurrently to events at different locations (e.g., the car in front, the pedestrian to the side). While spatial attention can be divided effectively between separate locations, studies investigating attention to nonspatial features have often reported a "global effect", whereby items having the attended feature may be preferentially processed throughout the entire visual field. These findings suggest that spatial and feature-based attention may at times act in direct opposition: spatially divided foci of attention cannot be truly independent if feature attention is spatially global and thereby affects all foci equally. In two experiments, human observers attended concurrently to one of two overlapping fields of dots of different colors presented in both the left and right visual fields. When the same color or two different colors were attended on the two sides, deviant targets were detected accurately, and visual-cortical potentials elicited by attended dots were enhanced. However, when the attended color on one side matched the ignored color on the opposite side, attentional modulation of cortical potentials was abolished. This loss of feature selectivity could be attributed to enhanced processing of unattended items that shared the color of the attended items in the opposite field. Thus, while it is possible to attend to two different colors at the same time, this ability is fundamentally constrained by spatially global feature enhancement in early visual-cortical areas, which is obligatory and persists even when it explicitly conflicts with task demands.

Salient sounds activate human visual cortex automatically.

Sudden changes in the acoustic environment enhance perceptual processing of subsequent visual stimuli that appear in close spatial proximity. Little is known, however, about the neural mechanisms by which salient sounds affect visual processing. In particular, it is unclear whether such sounds automatically activate visual cortex. To shed light on this issue, this study examined event-related brain potentials (ERPs) that were triggered either by peripheral sounds that preceded task-relevant visual targets (Experiment 1) or were presented during purely auditory tasks (Experiments 2-4). In all experiments the sounds elicited a contralateral ERP over the occipital scalp that was localized to neural generators in extrastriate visual cortex of the ventral occipital lobe. The amplitude of this cross-modal ERP was predictive of perceptual judgments about the contrast of colocalized visual targets. These findings demonstrate that sudden, intrusive sounds reflexively activate human visual cortex in a spatially specific manner, even during purely auditory tasks when the sounds are not relevant to the ongoing task.

Early auditory evoked potential is modulated by selective attention and related to individual differences in visual working memory capacity.

A growing body of research suggests that the predictive power of working memory (WM) capacity for measures of intellectual aptitude is due to the ability to control attention and select relevant information. Crucially, attentional mechanisms implicated in controlling access to WM are assumed to be domain-general, yet reports of enhanced attentional abilities in individuals with larger WM capacities are primarily within the visual domain. Here, we directly test the link between WM capacity and early attentional gating across sensory domains, hypothesizing that measures of visual WM capacity should predict an individual's capacity to allocate auditory selective attention. To address this question, auditory ERPs were recorded in a linguistic dichotic listening task, and individual differences in ERP modulations by attention were correlated with estimates of WM capacity obtained in a separate visual change detection task. Auditory selective attention enhanced ERP amplitudes at an early latency (ca. 70-90 msec), with larger P1 components elicited by linguistic probes embedded in an attended narrative. Moreover, this effect was associated with greater individual estimates of visual WM capacity. These findings support the view that domain-general attentional control mechanisms underlie the wide variation of WM capacity across individuals.

Spatio-temporal patterns of brain activity distinguish strategies of multiple-object tracking.

Human observers can readily track up to four independently moving items simultaneously, even in the presence of moving distractors. Here we combined EEG and magnetoencephalography recordings to investigate the neural processes underlying this remarkable capability. Participants were instructed to track four of eight independently moving items for 3 sec. When the movement ceased a probe stimulus consisting of four items with a higher luminance was presented. The location of the probe items could correspond fully, partly, or not at all with the tracked items. Participants reported whether the probe items fully matched the tracked items or not. About half of the participants showed slower RTs and higher error rates with increasing correspondence between tracked items and the probe. The other half, however, showed faster RTs and lower error rates when the probe fully matched the tracked items. This latter behavioral pattern was associated with enhanced probe-evoked neural activity that was localized to the lateral occipital cortex in the time range 170-210 msec. This enhanced response in the object-selective lateral occipital cortex suggested that these participants performed the tracking task by visualizing the overall shape configuration defined by the vertices of the tracked items, thereby producing a behavioral advantage on full-match trials. In a later time range (270-310 msec) probe-evoked neural activity increased monotonically as a function of decreasing target-probe correspondence in all participants. This later modulation, localized to superior parietal cortex, was proposed to reflect the degree of mismatch between the probe and the automatically formed visual STM representation of the tracked items.

Gamma band activity and the P3 reflect post-perceptual processes, not visual awareness.

A primary goal in cognitive neuroscience is to identify neural correlates of conscious perception (NCC). By contrasting conditions in which subjects are aware versus unaware of identical visual stimuli, a number of candidate NCCs have emerged; among them are induced gamma band activity in the EEG and the P3 event-related potential. In most previous studies, however, the critical stimuli were always directly relevant to the subjects' task, such that aware versus unaware contrasts may well have included differences in post-perceptual processing in addition to differences in conscious perception per se. Here, in a series of EEG experiments, visual awareness and task relevance were manipulated independently. Induced gamma activity and the P3 were absent for task-irrelevant stimuli regardless of whether subjects were aware of such stimuli. For task-relevant stimuli, gamma and the P3 were robust and dissociable, indicating that each reflects distinct post-perceptual processes necessary for carrying-out the task but not for consciously perceiving the stimuli. Overall, this pattern of results challenges a number of previous proposals linking gamma band activity and the P3 to conscious perception.

Audio-visual synchrony modulates the ventriloquist illusion and its neural/spatial representation in the auditory cortex.

An essential task of our perceptual systems is to bind together the distinctive features of single objects and events into unitary percepts, even when those features are registered in different sensory modalities. In cases where auditory and visual inputs are spatially incongruent, they may still be perceived as belonging to a single event at the location of the visual stimulus - a phenomenon known as the 'ventriloquist illusion'. The present study examined how audio-visual temporal congruence influences the ventriloquist illusion and characterized its neural underpinnings with functional magnetic resonance imaging (fMRI). Behaviorally, the ventriloquist illusion was reduced for asynchronous versus synchronous audio-visual stimuli, in accordance with previous reports. Neural activity patterns associated with the ventriloquist effect were consistently observed in the planum temporale (PT), with a reduction in illusion-related fMRI-signals ipsilateral to visual stimulation for central sounds perceived peripherally and a contralateral increase in illusion-related fMRI-signals for peripheral sounds perceived centrally. Moreover, it was found that separate but adjacent regions within the PT were preferentially activated for ventriloquist illusions produced by synchronous and asynchronous audio-visual stimulation. We conclude that the left-right balance of neural activity in the PT represents the neural code that underlies the ventriloquist illusion, with greater activity in the cerebral hemisphere contralateral to the direction of the perceived shift of sound location.

Earliest stages of visual cortical processing are not modified by attentional load.

This study investigated the effects of attentional load on neural responses to attended and irrelevant visual stimuli by recording high-density event-related potentials (ERPs) from the scalp in normal adult subjects. Peripheral (upper and lower visual field) and central stimuli were presented in random order at a rapid rate while subjects responded to targets among the central stimuli. Color detection and color-orientation conjunction search tasks were used as the low- and high-load tasks, respectively. Behavioral results showed significant load effects on both accuracy and reaction time for target detections. ERP results revealed no significant load effect on the initial C1 component (60-100 ms) evoked by either central-relevant or peripheral-irrelevant stimuli. Source analysis with dipole modeling confirmed previous reports that the C1 includes the initial evoked response in primary visual cortex. Source analyses indicated that high attentional load enhanced the early (70-140 ms) neural response to central-relevant stimuli in ventral-lateral extrastriate cortex, whereas load effects on peripheral-irrelevant stimulus processing started at 110 ms and were localized to more dorsal and anterior extrastriate cortical areas. These results provide evidence that the earliest stages of visual cortical processing are not modified by attentional load and show that attentional load affects the processing of task relevant and irrelevant stimuli in different ways.

Parietal connectivity mediates multisensory facilitation.

Our senses interact in daily life through multisensory integration, facilitating perceptual processes and behavioral responses. The neural mechanisms proposed to underlie this multisensory facilitation include anatomical connections directly linking early sensory areas, indirect connections to higher-order multisensory regions, as well as thalamic connections. Here we examine the relationship between white matter connectivity, as assessed with diffusion tensor imaging, and individual differences in multisensory facilitation and provide the first demonstration of a relationship between anatomical connectivity and multisensory processing in typically developed individuals. Using a whole-brain analysis and contrasting anatomical models of multisensory processing we found that increased connectivity between parietal regions and early sensory areas was associated with the facilitation of reaction times to multisensory (auditory-visual) stimuli. Furthermore, building on prior animal work suggesting the involvement of the superior colliculus in this process, using probabilistic tractography we determined that the strongest cortical projection area connected with the superior colliculus includes the region of connectivity implicated in our independent whole-brain analysis.

Consequences of magnocellular dysfunction on processing attended information in schizophrenia.

Schizophrenia is associated with perceptual and cognitive dysfunction including impairments in visual attention. These impairments may be related to deficits in early stages of sensory/perceptual processing, particularly within the magnocellular/dorsal visual pathway. In the present study, subjects viewed high and low spatial frequency (SF) gratings designed to test functioning of the parvocellular/magnocellular pathways, respectively. Schizophrenia patients and healthy controls attended to either the low SF (magnocellularly biased) or high SF (parvocellularly biased) gratings. Functional magnetic resonance imaging (fMRI) and recordings of event-related potentials (ERPs) were carried out during task performance. Patients were impaired at detecting low-frequency targets. ERP amplitudes to low-frequency gratings were diminished, both for the early sensory-evoked components and for the attend minus unattend difference component (the selection negativity), which is regarded as a neural index of feature-selective attention. Similarly, fMRI revealed that activity in extrastriate visual cortex was reduced in patients during attention to low, but not high, SF. In contrast, activity in frontal and parietal areas, previously implicated in the control of attention, did not differ between patients and controls. These findings suggest that impaired sensory processing of magnocellularly biased stimuli lead to impairments in the effective processing of attended stimuli, even when the attention control systems themselves are intact.