In an explorative study, we investigated the time course of attentional selection shifts in feature-based attention in early visual cortex by means of steady-state visual evoked potentials (SSVEPs). To this end, we presented four flickering random dot kinematograms with red/blue, horizontal/vertical bars, respectively. Given the oscillatory nature of SSVEPs, we were able to investigate neural temporal dynamics of facilitation and inhibition/suppression when participants shifted attention either within (i.e., color to color) or between feature dimensions (i.e., color to orientation). Extending a previous study of our laboratory [Müller, M. M., Trautmann, M., & Keitel, C. Early visual cortex dynamics during top–down modulated shifts of feature-selective attention. Journal of Cognitive Neuroscience , 28 , 643–655, 2016] to a full factorial design, we replicated a critical finding of our previous study: Facilitation of color was quickest, regardless of the origin of the shift (from color or orientation). Furthermore, facilitation of the newly to-be-attended and inhibition/suppression of the then to-be-ignored feature is not a time-invariant process that occurs instantaneously, but a biphasic one with longer time delays between the two processes. Interestingly, inhibition/suppression of the to-be-ignored feature after the shifting cue had a much longer latency with between- compared to within-dimensional shifts (by about 130–150 msec). The exploratory nature of our study is reasoned by two limiting factors: (a) Identical to our precursor study, we found no attentional SSVEP amplitude time course modulation for orientation, and (b) the signal-to-noise ratio for single trials was too poor to allow for reliable statistical testing of the latencies that were obtained with running t tests of averaged data.

We assessed the extent of neural competition for attentional processing resources in early visual cortex between foveally presented task stimuli and peripheral emotional distracter images. Task-relevant and distracting stimuli were shown in rapid serial visual presentation (RSVP) streams to elicit the steady-state visual evoked potential, which serves as an electrophysiological marker of attentional resource allocation in early visual cortex. A task-related RSVP stream of symbolic letters was presented centrally at 15 Hz while distracting RSVP streams were displayed at 4 or 6 Hz in the left and right visual hemifields. These image streams always had neutral content in one visual field and would unpredictably switch from neutral to unpleasant content in the opposite visual field. We found that the steady-state visual evoked potential amplitude was consistently modulated as a function of change in emotional valence in peripheral RSVPs, indicating sensory gain in response to distracting affective content. Importantly, the facilitated processing for emotional content shown in one visual hemifield was not paralleled by any perceptual costs in response to the task-related processing in the center or the neutral image stream in the other visual hemifield. Together, our data provide further evidence for sustained sensory facilitation in favor of emotional distracters. Furthermore, these results are in line with previous reports of a “different hemifield advantage” with low-level visual stimuli and are suggestive of independent processing resources in each cortical hemisphere that operate beyond low-level visual cues, that is, with complex images that impact early stages of visual processing via reentrant feedback loops from higher order processing areas.

The integrated object account predicts that attention is spread across all features that constitute one object, regardless of their task relevance. We challenge that prediction with a novel stimulation technique that allows for simultaneous electrophysiological measurements of the allocation of attention to two distinct features within one object. A rotating square that flickers in different colors evoked two distinct steady-state visual evoked potentials (SSVEPs) for rotation and color, respectively. If the integrated object account were true, we would expect identical SSVEP amplitudes regardless of what feature participants attended. We found greater SSVEP amplitudes for the to-be-attended feature compared with the to-be-ignored feature. SSVEP amplitudes averaged across both features were significantly reduced when participants attended to both features, which was mirrored in behavioral costs, implying competitive interactions or a division of attentional resources. Surprisingly, this reduction in amplitude was mainly driven by the SSVEP amplitude elicited by color changes. In conclusion, our results challenge the integrated object account and highlight the extent to which color is “special” within feature space.

A key property of feature-based attention is global facilitation of the attended feature throughout the visual field. Previously, we presented superimposed red and blue randomly moving dot kinematograms (RDKs) flickering at a different frequency each to elicit frequency-specific steady-state visual evoked potentials (SSVEPs) that allowed us to analyze neural dynamics in early visual cortex when participants shifted attention to one of the two colors. Results showed amplification of the attended and suppression of the unattended color as measured by SSVEP amplitudes. Here, we tested whether the suppression of the unattended color also operates globally. To this end, we presented superimposed flickering red and blue RDKs in the center of a screen and a red and blue RDK in the left and right periphery, respectively, also flickering at different frequencies. Participants shifted attention to one color of the superimposed RDKs in the center to discriminate coherent motion events in the attended from the unattended color RDK, whereas the peripheral RDKs were task irrelevant. SSVEP amplitudes elicited by the centrally presented RDKs confirmed the previous findings of amplification and suppression. For peripherally located RDKs, we found the expected SSVEP amplitude increase, relative to precue baseline when color matched the one of the centrally attended RDK. We found no reduction in SSVEP amplitude relative to precue baseline, when the peripheral color matched the unattended one of the central RDK, indicating that, while facilitation in feature-based attention operates globally, suppression seems to be linked to the location of focused attention.

Shifting attention from one color to another color or from color to another feature dimension such as shape or orientation is imperative when searching for a certain object in a cluttered scene. Most attention models that emphasize feature-based selection implicitly assume that all shifts in feature-selective attention underlie identical temporal dynamics. Here, we recorded time courses of behavioral data and steady-state visual evoked potentials (SSVEPs), an objective electrophysiological measure of neural dynamics in early visual cortex to investigate temporal dynamics when participants shifted attention from color or orientation toward color or orientation, respectively. SSVEPs were elicited by four random dot kinematograms that flickered at different frequencies. Each random dot kinematogram was composed of dashes that uniquely combined two features from the dimensions color (red or blue) and orientation (slash or backslash). Participants were cued to attend to one feature (such as color or orientation) and respond to coherent motion targets of the to-be-attended feature. We found that shifts toward color occurred earlier after the shifting cue compared with shifts toward orientation, regardless of the original feature (i.e., color or orientation). This was paralleled in SSVEP amplitude modulations as well as in the time course of behavioral data. Overall, our results suggest different neural dynamics during shifts of attention from color and orientation and the respective shifting destinations, namely, either toward color or toward orientation.

Visual attention can be focused concurrently on two stimuli at noncontiguous locations while intermediate stimuli remain ignored. Nevertheless, behavioral performance in multifocal attention tasks falters when attended stimuli fall within one visual hemifield as opposed to when they are distributed across left and right hemifields. This “different-hemifield advantage” has been ascribed to largely independent processing capacities of each cerebral hemisphere in early visual cortices. Here, we investigated how this advantage influences the sustained division of spatial attention. We presented six isoeccentric light-emitting diodes (LEDs) in the lower visual field, each flickering at a different frequency. Participants attended to two LEDs that were spatially separated by an intermediate LED and responded to synchronous events at to-be-attended LEDs. Task-relevant pairs of LEDs were either located in the same hemifield (“within-hemifield” conditions) or separated by the vertical meridian (“across-hemifield” conditions). Flicker-driven brain oscillations, steady-state visual evoked potentials (SSVEPs), indexed the allocation of attention to individual LEDs. Both behavioral performance and SSVEPs indicated enhanced processing of attended LED pairs during “across-hemifield” relative to “within-hemifield” conditions. Moreover, SSVEPs demonstrated effective filtering of intermediate stimuli in “across-hemifield” condition only. Thus, despite identical physical distances between LEDs of attended pairs, the spatial profiles of gain effects differed profoundly between “across-hemifield” and “within-hemifield” conditions. These findings corroborate that early cortical visual processing stages rely on hemisphere-specific processing capacities and highlight their limiting role in the concurrent allocation of visual attention to multiple locations.

The focus of attention can be flexibly altered in mnemonic representations of past sensory events. We investigated the neural mechanisms of selection in tactile STM by applying vibrotactile sample stimuli of different intensities to both hands, followed by a symmetrically shaped visual retro-cue. The retro-cue indicated whether the weak or strong sample was relevant for subsequent comparison with a single tactile test stimulus. Locations of tactile stimuli were randomized, and the required response did not depend upon the spatial relation between cued sample and test stimulus. Selection between spatially segregated items in tactile STM was mirrored in lateralized activity following visual retro-cues (N2pc) and influenced encoding of task-irrelevant tactile probe stimuli (N140). Our findings support four major conclusions. First, retrospective selection results in transient shifts of spatial attention. Second, retrospective selection is functionally dissociable from attention-based rehearsal of locations. Third, selection mechanisms are linked across processing stages, as attention shifts in STM influence encoding of sensory signals. Fourth, selection in tactile STM recruits attentional control mechanisms that are, at least partially, supramodal.

We investigated mechanisms of concurrent attentional selection of location and color using electrophysiological measures in human subjects. Two completely overlapping random dot kinematograms (RDKs) of two different colors were presented on either side of a central fixation cross. On each trial, participants attended one of these four RDKs, defined by its specific combination of color and location, in order to detect coherent motion targets. Sustained attentional selection while monitoring for targets was measured by means of steady-state visual evoked potentials (SSVEPs) elicited by the frequency-tagged RDKs. Attentional selection of transient targets and distractors was assessed by behavioral responses and by recording event-related potentials to these stimuli. Spatial attention and attention to color had independent and largely additive effects on the amplitudes of SSVEPs elicited in early visual areas. In contrast, behavioral false alarms and feature-selective modulation of P3 amplitudes to targets and distractors were limited to the attended location. These results suggest that feature-selective attention produces an early, global facilitation of stimuli having the attended feature throughout the visual field, whereas the discrimination of target events takes place at a later stage of processing that is only applied to stimuli at the attended position.

Object recognition is achieved through neural mechanisms reliant on the activity of distributed neural assemblies that are thought to be coordinated by synchronous firing in the gamma-band range (>20 Hz). An outstanding question focuses on the extent to which the role of gamma oscillations in object recognition is dependent on attention. Attentional mechanisms determine the allocation of perceptual resources to objects in complex scenes biasing the outcome of their mutual competitive interactions. Would object-related enhancements in gamma activity also occur for unattended objects when perceptual resources are traded off to the processing of concurrent visual material? The present electroencephalogram study investigated event-related potentials and evoked (time- and phase-locked) and induced (non-time- and phase-locked to stimulus onset) gamma-band activity (GBA) using a visual discrimination task of low or high perceptual load at fixation. The task was performed while task-irrelevant familiar or unfamiliar objects coappeared in the surrounding central area. Attentional focus was kept at fixation by varying perceptual load between trials; in such conditions, only holistic object processing or low-level perceptual processing, requiring little or no attention, are thought to occur. Although evoked GBA remained unmodulated, induced GBA enhancements, specific to familiar object presentations, were observed, thus providing evidence for cortical visual representation of unattended objects. In addition, the effect was mostly driven by object-specific activity under low load, implying that, in cluttered or complex scenes, attentional selection likely plays a more significant role in object representation.

Modulations of oscillatory electroencephalogram (EEG) activity in the induced gamma and theta frequency ranges (induced gamma and theta band responses; iGBRs: >30 Hz; iTBRs: ∼6 Hz) have been associated with retrieval of information from long-term memory. However, the specific functional role of these two forms of oscillatory activity remains unclear. The present study examines theta- and gamma-oscillations within a dual-process framework, which defines “familiarity” and “recollection” as the two component processes of recognition memory. During encoding, participants were instructed to make “bigger/smaller than a shoebox” or “living/nonliving” decisions for different object pictures. During retrieval “old/new” recognition was followed (for items judged old) by a source discrimination task regarding the decision made for each item at encoding. iGBRs (35–80 Hz; 210–330 msec) were higher for correctly identified “old” relative to “new” objects. Importantly, they did not distinguish between successful and unsuccessful source judgments. In contrast, iTBRs (4–7.5 Hz; 600–1200 msec) were sensitive to source discrimination. We propose that iGBRs mirror early associative processes linked to familiarity-related retrieval processes, whereas iTBRs reflect later onsetting, episodic, recollection-related mechanisms.

Neural mechanisms of object recognition seem to rely on activity of distributed neural assemblies coordinated by synchronous firing in the gamma-band range (>20 Hz). In the present electroencephalogram (EEG) study, we investigated induced gamma band activity during the naming of line drawings of upright objects and objects rotated in the image plane. Such plane-rotation paradigms elicit view-dependent processing, leading to delays in recognition of disoriented objects. Our behavioral results showed reaction time delays for rotated, as opposed to upright, images. These delays were accompanied by delays in the peak latency of induced gamma band responses (GBRs), in the absence of any effects on other measures of EEG activity. The latency of the induced GBRs has thus, for the first time, been selectively modulated by an experimental manipulation that delayed recognition. This finding indicates that induced GBRs have a genuine role as neural markers of late representational processes during object recognition. In concordance with the view that object recognition is achieved through dynamic learning processes, we propose that induced gamma band activity could be one of the possible cortical markers of such dynamic object coding.

In the present study, subjects selectively attended to the color of checkerboards in a feature-based attention paradigm. Induced gamma band responses (GBRs), the induced alpha band, and the event-related potential (ERP) were analyzed to uncover neuronal dynamics during selective feature processing. Replicating previous ERP findings, the selection negativity (SN) with a latency of about 160 msec was extracted. Furthermore, and similarly to previous EEG studies, a gamma band peak in a time window between 290 and 380 msec was found. This peak had its major energy in the 55to 70-Hz range and was significantly larger for the attended color. Contrary to previous human induced gamma band studies, a much earlier 40to 50-Hz peak in a time window between 160 and 220 msec after stimulus onset and, thus, concurrently to the SN was prominent with significantly more energy for attended as opposed to unattended color. The induced alpha band (9.8–11.7 Hz), on the other hand, exhibited a marked suppression for attended color in a time window between 450 and 600 msec after stimulus onset. A comparison of the time course of the 40to 50-Hz and 55to 70-Hz induced GBR, the induced alpha band, and the ERP revealed temporal coincidences for changes in the morphology of these brain responses. Despite these similarities in the time domain, the cortical source configuration was found to discriminate between induced GBRs and the SN. Our results suggest that large-scale synchronous high-frequency brain activity as measured in the human GBR play a specific role in attentive processing of stimulus features.

Fragmented pictures of an object, which appear meaningless when seen for the first time, can easily be identified after the presentation of an unfragmented version of the same picture. The neuronal mechanism for such a rapid perceptual learning phenomenon is largely unknown. Recently, induced gamma band responses (GBRs) have been discussed as a possible physiological correlate of activity in cell assemblies formed by learning. The present study was designed to investigate the modulation of induced GBRs in a perceptual learning task by using a 128-channel EEG montage. In the first sequence of the experiment, fragmented pictures from the Snodgrass and Vandervart inventory were presented. The fragmentation of the pictures was selected that subjects were unable to identify them. In the second experimental sequence the perceptual learning sequence half of the pictures were displayed in their unfragmented version. In the third sequence, all pictures were presented again in the fragmented version. Now, subjects had to rate whether or not they could identify the images. Results showed an increase in spectral gamma power at parietal electrode sites for identified pictures. In addition, neural activity in the gamma band was highly synchronized between posterior electrodes. For pictures not presented in their complete version, we found no such pattern in the third sequence. From our results, we concluded that induced GBRs might represent a signature of synchronized neural activity in a Hebbian cell assembly, activated by the fragmented picture after perceptual learning took place. No difference between identified and unidentified pictures was found in the visual evoked potential in the same time range and in the evoked GBR in the same frequency range as the induced response.