Distinct patterns of spatial attentional modulation of steady-state visual evoked magnetic fields (SSVEFs) in subdivisions of the human early visual cortex.

In recent years, steady-state visual evoked potentials (SSVEPs) became an increasingly valuable tool to investigate neural dynamics of competitive attentional interactions and brain-computer interfaces. This is due to their good signal-to-noise ratio, allowing for single-trial analysis, and their ongoing oscillating nature that enables to analyze temporal dynamics of facilitation and suppression. Given the popularity of SSVEPs, it is surprising that only a few studies looked at the cortical sources of these responses. This is in particular the case when searching for studies that assessed the cortical sources of attentional SSVEP amplitude modulations. To address this issue, we used a typical spatial attention task and recorded neuromagnetic fields (MEG) while presenting frequency-tagged stimuli in the left and right visual fields, respectively. Importantly, we controlled for attentional deployment in a baseline period before the shifting cue. Subjects either attended to a central fixation cross or to two peripheral stimuli simultaneously. Results clearly showed that signal sources and attention effects were restricted to the early visual cortex: V1, V2, hMT+, precuneus, occipital-parietal, and inferior-temporal cortex. When subjects attended to central fixation first, shifting attention to one of the peripheral stimuli resulted in a significant activation increase for the to-be-attended stimulus with no activation decrease for the to-be-ignored stimulus in hMT+ and inferio-temporal cortex, but significant SSVEF decreases from V1 to occipito-parietal cortex. When attention was first deployed to both rings, shifting attention away from one ring basically resulted in a significant activation decrease in all areas for the then-to-be-ignored stimulus.

Early sensory gain control is dominated by obligatory and global feature-based attention in top-down shifts of combined spatial and feature-based attention.

What are the dynamics of global feature-based and spatial attention, when deployed together? In an attentional shifting experiment, flanked by three control experiments, we investigated neural temporal dynamics of combined attentional shifts. For this purpose, orange- and blue-frequency-tagged spatially overlapping Random Dot Kinematograms were presented in the left and right visual hemifield to elicit continuous steady-state-visual-evoked-potentials. After being initially engaged in a fixation cross task, participants were at some point in time cued to shift attention to one of the Random Dot Kinematograms, to detect and respond to brief coherent motion events, while ignoring all such events in other Random Dot Kinematograms. The analysis of steady-state visual-evoked potentials allowed us to map time courses and dynamics of early sensory-gain modulations by attention. This revealed a time-invariant amplification of the to-be attended color both at the attended and the unattended side, followed by suppression for the to-be-ignored color at attended and unattended sides. Across all experiments, global and obligatory feature-based selection dominated early sensory gain modulations, whereas spatial attention played a minor modulatory role. However, analyses of behavior and neural markers such as alpha-band activity and event-related potentials to target- and distractor-event processing, revealed clear modulations by spatial attention.

Global enhancement of target color-not proactive suppression-explains attentional deployment during visual search.

The current study touches on a central debate in the area of attention: how the human brain handles distraction by salient stimuli. The idea of proactive suppression proposes a new fundamental perceptual mechanism to resolve this question, whereby attentional capture by a task-irrelevant salient distractor can be preempted through top-down inhibitory mechanisms. In this study, we replicate empirical effects underlying this claim, but show that they are better explained by an alternative mechanism, global target-feature enhancement. Identical to original studies using a capture-probe dual-task design, observers recalled fewer letters superimposed upon color singleton distractors, relative to other irrelevant search items (fillers). However, given that fillers (but not singleton distractors) always matched the color of the target, this effect could have been due to global featural attention to the target color rather than suppression of the singleton distractor. After manipulating the color of fillers such that they no longer matched the target color, probe recall associated with these was reduced, causing the relative "suppression" of singleton distractors to be abolished. We then manipulated the color similarity of targets and fillers, and found that filler probe recall was graded as a function of this color similarity, even within a single search context. This strongly suggests that increased attention to fillers due to global target color enhancement underlies the difference in attention among distractor items, not proactive distractor suppression. In contrast with feature enhancement and reactive suppression, the proposed proactive suppression mechanism still lacks convincing behavioral evidence. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Higher amplitudes in steady-state visual evoked potentials driven by square-wave versus sine-wave contrast modulation - A dual-laboratory study.

Steady-state visual evoked potentials (ssVEPs) are an established tool for assessing visuocortical responses in visual perception and attention. They have the same temporal frequency characteristics as a periodically modulated stimulus (e.g., in contrast or luminance) that drives them. It has been hypothesized that the amplitude of a given ssVEP may depend on the shape of the stimulus modulation function, but the size and robustness of these effects is not well established. The current study systematically compared the effect of the two most common functions in the ssVEP literature, square-wave and sine-wave functions. Across two laboratories, we presented mid-complex color patterns to 30 participants with square-wave or sine-wave contrast modulation and at different driving frequencies (6 Hz, 8.57 Hz, 15 Hz). When ssVEPs were analyzed independently for the samples, with each laboratory's standard processing pipeline, ssVEP amplitudes in both samples decreased at higher driving frequencies and square-wave modulation evoked higher amplitudes at lower frequencies (i.e., 6 Hz, 8.57 Hz) compared to sine-wave modulation. These effects were replicated when samples were aggregated and analyzed with the same processing pipeline. In addition, when using signal-to-noise ratios as outcome measures, this joint analysis indicated a somewhat weaker effect of increased ssVEP amplitudes to square-wave modulation at 15 Hz. The present study suggests that square-wave modulation should be used in ssVEP research when the goal is to maximize signal amplitude or signal-to-noise ratio. Given effects of modulation function across laboratories, and data processing pipelines, the findings appear robust to differences in data collection and analysis.

Global attentional selection of visual features is not associated with selective modulation of posterior alpha-band activity.

Attending to a single feature, such as color or motion, leads to global modulation of neural processing associated with the representation of the attended features. Alpha-band modulations are hypothesized to be a marker (and even a mechanism) of the modulation of neural processing. By adopting a previously used attentional shifting paradigm, we examined whether alpha-band dynamics are linked to sustained Feature-Based-Attentional (FBA) selection. For this purpose, we presented task-irrelevant flickering random dot kinematograms (RDKs) in the periphery that either did or did not share the to-be-attended color of centrally presented task-relevant RDKs and should thus be subject to global FBA selection. Steady-state visual evoked potentials (SSVEPs) and alpha-band activity associated with these task-irrelevant RDKs were analyzed to quantify FBA modulation. Overall, the SSVEP results replicated previous findings: relative to a pre-cue baseline, SSVEP amplitudes for peripheral RDKs were significantly enhanced when these RDKs shared the to-be-attended color of the central RDKs and were not modulated when they shared the centrally to-be-ignored color. Nevertheless, there were no differences in alpha-band amplitude modulations between signals recorded contralateral to the RDKs sharing the centrally attended color and RDKs sharing the centrally ignored color. Hence, alpha-band modulations seem not to index the sustained global selection of attended over unattended feature values within the same feature dimension.

Attentional capture is modulated by stimulus saliency in visual search as evidenced by event-related potentials and alpha oscillations.

This study used a typical four-item search display to investigate top-down control over attentional capture in an additional singleton paradigm. By manipulating target and distractor color and shape, stimulus saliency relative to the remaining items was systematically varied. One group of participants discriminated the side of a dot within a salient orange target (ST group) presented with green circles (fillers) and a green diamond distractor. A second group discriminated the side of the dot within a green diamond target presented with green circle fillers and a salient orange square distractor (SD group). Results showed faster reaction times and a shorter latency of the N2pc component in the event-related potential (ERP) to the more salient targets in the ST group. Both salient and less salient distractors elicited Pd components of equal amplitude. Behaviorally, no task interference was observed with the less salient distractor, indicating the prevention of attentional capture. However, reaction times were slower in the presence of the salient distractor, which conflicts with the hypothesis that the Pd reflects proactive distractor suppression. Contrary to recent proposals that elicitation of the Pd requires competitive interactions with a target, we found a greater Pd amplitude when the distractor was presented alone. Alpha-band amplitudes decreased during target processing (event-related desynchronization), but no significant amplitude enhancement was observed at electrodes contralateral to distractors regardless of their saliency. The results demonstrate independent neural mechanisms for target and distractor processing and support the view that top-down guidance of attention can be offset (counteracted) by relative stimulus saliency.

Dynamics of attentional allocation to targets and distractors during visual search.

There is much debate about the neural mechanisms that achieve suppression of salient distracting stimuli during visual search. The proactive suppression hypothesis asserts that if exposed to the same distractors repeatedly, these stimuli are actively inhibited before attention can be shifted to them. A contrasting proposal holds that attention is initially captured by salient distractors but is subsequently withdrawn. By concurrently measuring stimulus-driven and intrinsic brain potentials in 36 healthy human participants, we obtained converging evidence against early proactive suppression of distracting input. Salient distractors triggered negative event-related potentials (N1pc/N2pc), enhanced the steady-state visual evoked potential (SSVEP) relative to non-salient (filler) stimuli, and suppressed contralateral relative to ipsilateral alpha-band amplitudes-three electrophysiological measure associated with the allocation of attention-even though these distractors did not interfere with behavioral responses to the search targets. Furthermore, these measures indicated that both stimulus-driven and goal-driven allocations of attention occurred in conjunction with one another, with the goal-driven effect enhancing and prolonging the stimulus-driven effect. These results provide a new perspective on the traditional dichotomy between bottom-up and top-down attentional allocation. Control experiments revealed that continuous marking of the locations at which the search display items were presented resulted in a dramatic and unexpected conversion of the target-elicited N2pc into a shorter-latency N1pc in association with faster reaction times to the targets.

Electrophysiological evidence for target facilitation without distractor suppression in two-stimulus search displays.

This study used electrophysiological measures to investigate how attention is deployed to target and distractor stimuli during visual search using search displays with a small set-size. Participants viewed randomized sequences of two-item displays that consisted of either a target and a distractor (differing in color) or a pair of task-irrelevant filler stimuli having a third color, all presented in an ongoing stream of flickering gray circles. The allocation of attention was assessed by concurrent recordings of steady-state visual evoked potentials (SSVEPs) elicited by the flickering displays and perturbations of the endogenous alpha rhythm following each type of display. The aim was to test a central prediction of the signal suppression hypothesis, namely that the processing of distractors will be proactively suppressed below the level of filler stimuli. Amplitude modulations of both the SSVEP and the lateralized alpha rhythm provided converging evidence against early proactive suppression of highly salient distractors. Instead, these electrophysiological measures were consistent with the view that in this type of two-stimulus search task there is an initial capture of attention by all color-change stimuli (targets, distractors, and fillers) followed by a further focusing of attention upon the target, with no evidence for suppression of the distractor.

Suppression of Unattended Features Is Independent of Task Relevance.

Feature-based attention serves the separation of relevant from irrelevant features. While global amplification of attended features is coherently described as a key mechanism for feature-based attention, nature and constituting factors of neural suppressive interactions are far less clear. One aspect of global amplification is its flexible modulation by the task relevance of the to-be-attended stimulus. We examined whether suppression is similarly modulated by their respective task relevance or is mandatory for all unattended features. For this purpose, participants saw a display of randomly moving dots with 3 distinct colors and were asked to report brief events of coherent motion for a cued color. Of the 2 unattended colored clouds, one contained distracting motion events while the other was irrelevant and without such motion events throughout the experiment. We used electroencephalography-derived steady-state visual-evoked potentials to investigate early visual processing of the attended, unattended, and irrelevant color under sustained feature-based attention. The analysis revealed a biphasic process with an early amplification of the to-be-attended color followed by suppression of the to-be-ignored color relative to a pre-cue baseline. Importantly, the neural dynamics for the unattended and always irrelevant color were comparable. Suppression is thus a mandatory mechanism affecting all unattended stimuli irrespective of their task relevance.

Alpha and beta neural oscillations differentially reflect age-related differences in bilateral coordination.

Bilateral in-phase (IP) and anti-phase (AP) movements represent two fundamental modes of bilateral coordination that are essential for daily living. Although previous studies have shown that aging is behaviorally associated with decline in bilateral coordination, especially in AP movements, the underlying neural mechanisms remain unclear. Here, we use kinematic measurements and electroencephalography to compare motor performance of young and older adults executing bilateral IP and AP hand movements. On the behavioral level, inter-limb synchronization was reduced during AP movements compared to IP and this reduction was stronger in the older adults. On the neural level, we found interactions between group and condition for task-related power change in different frequency bands. The interaction was driven by smaller alpha power decreases over the non-dominant cortical motor area in young adults during IP movements and larger beta power decreases over the midline region in older adults during AP movements. In addition, the decrease in inter-limb synchronization during AP movements was predicted by stronger directional connectivity in the beta-band: an effect more pronounced in older adults. Our results therefore show that age-related differences in the two bilateral coordination modes are reflected on the neural level by differences in alpha and beta oscillatory power as well as interhemispheric directional connectivity.

Transient Amplitude Modulation of Alpha-Band Oscillations by Short-Time Intermittent Closed-Loop tACS.

Non-invasive brain stimulation (NIBS) techniques such as transcranial alternating current stimulation (tACS) have recently become extensively utilized due to their potential to modulate ongoing neuronal oscillatory activity and consequently to induce cortical plasticity relevant for various cognitive functions. However, the neurophysiological basis for stimulation effects as well as their inter-individual differences is not yet understood. In the present study, we used a closed-loop electroencephalography-tACS(EEG-tACS) protocol to examine the modulation of alpha oscillations generated in occipito-parietal areas. In particular, we investigated the effects of a repeated short-time intermittent stimulation protocol (1 s in every trial) applied over the visual cortex (Cz and Oz) and adjusted according to the phase and frequency of visual alpha oscillations on the amplitude of these oscillations. Based on previous findings, we expected higher increases in alpha amplitudes for tACS applied in-phase with ongoing oscillations as compared to an application in anti-phase and this modulation to be present in low-alpha amplitude states of the visual system (eyes opened, EO) but not high (eyes closed, EC). Contrary to our expectations, we found a transient suppression of alpha power in inter-individually derived spatially specific parieto-occipital components obtained via the estimation of spatial filters by using the common spatial patterns approach. The amplitude modulation was independent of the phase relationship between the tACS signal and alpha oscillations, and the state of the visual system manipulated via closed- and open-eye conditions. It was also absent in conventionally analyzed single-channel and multi-channel data from an average parieto-occipital region. The fact that the tACS modulation of oscillations was phase-independent suggests that mechanisms driving the effects of tACS may not be explained by entrainment alone, but rather require neuroplastic changes or transient disruption of neural oscillations. Our study also supports the notion that the response to tACS is subject-specific, where the modulatory effects are shaped by the interplay between the stimulation and different alpha generators. This favors stimulation protocols as well as analysis regimes exploiting inter-individual differences, such as spatial filters to reveal otherwise hidden stimulation effects and, thereby, comprehensively induce and study the effects and underlying mechanisms of tACS.

Reduction of somatosensory functional connectivity by transcranial alternating current stimulation at endogenous mu-frequency.

Alpha, the most prominent human brain rhythm, might reflect a mechanism of functional inhibition for gating neural processing. This concept has been derived predominantly from local measures of inhibition, while large-scale network mechanisms to guide information flow are largely unknown. Here, we investigated functional connectivity changes on a whole-brain level by concurrent transcranial alternating current stimulation (tACS) and resting-state functional MRI in humans. We specifically focused on somatosensory alpha-band oscillations by adjusting the tACS frequency to each individual´s somatosensory (mu-) alpha peak frequency (mu-tACS). Potential differences of Eigenvector Centrality of primary somatosensory cortex (S1) as well as on a whole brain level between mu-tACS and sham were analyzed. Our results demonstrate that mu-tACS induces a locally-specific decrease in whole-brain functional connectivity of left S1. An additional exploratory analysis revealed that this effect primarily depends on a decrease in functional connectivity between S1 and a network of regions that are crucially involved in somatosensory processing. Furthermore, the decrease in functional centrality was specific to mu-tACS and was not observed when tACS was applied in the gamma-range in an independent study. Our findings provide evidence that modulated somatosensory (mu-) alpha-activity may affect whole-brain network level activity by decoupling primary sensory areas from other hubs involved in sensory processing.

Rapid processing of neutral and angry expressions within ongoing facial stimulus streams: Is it all about isolated facial features?

Our visual system extracts the emotional meaning of human facial expressions rapidly and automatically. Novel paradigms using fast periodic stimulations have provided insights into the electrophysiological processes underlying emotional content extraction: the regular occurrence of specific identities and/or emotional expressions alone can drive diagnostic brain responses. Consistent with a processing advantage for social cues of threat, we expected angry facial expressions to drive larger responses than neutral expressions. In a series of four EEG experiments, we studied the potential boundary conditions of such an effect: (i) we piloted emotional cue extraction using 9 facial identities and a fast presentation rate of 15 Hz (N = 16); (ii) we reduced the facial identities from 9 to 2, to assess whether (low or high) variability across emotional expressions would modulate brain responses (N = 16); (iii) we slowed the presentation rate from 15 Hz to 6 Hz (N = 31), the optimal presentation rate for facial feature extraction; (iv) we tested whether passive viewing instead of a concurrent task at fixation would play a role (N = 30). We consistently observed neural responses reflecting the rate of regularly presented emotional expressions (5 Hz and 2 Hz at presentation rates of 15 Hz and 6 Hz, respectively). Intriguingly, neutral expressions consistently produced stronger responses than angry expressions, contrary to the predicted processing advantage for threat-related stimuli. Our findings highlight the influence of physical differences across facial identities and emotional expressions.

Rapid sensory gain with emotional distracters precedes attentional deployment from a foreground task.

The steady-state visual evoked potential (SSVEP), an electrophysiological marker of attentional resource allocation, has recently been demonstrated to serve as a neural signature of emotional content extraction from a rapid serial visual presentation (RSVP). SSVEP amplitude was reduced for streams of emotional relative to neutral scenes passively viewed at 6 Hz (~167 ms per image), but it was enhanced for emotional relative to neutral scenes when viewed as 4 Hz RSVP (250 ms per image). Here, we investigated whether these seemingly contradictory observations may be related to different dynamics in the allocation of attentional resources as a consequence of stimulation frequency. To this end, we advanced our distraction paradigm by presenting a visual foreground task consisting of randomly moving squares flickering at 15 Hz superimposed on task-irrelevant RSVP streams shown at 6 or 4 Hz, which could unpredictably switch from neutral to unpleasant content during the trial or remained neutral. Critically, our findings demonstrate that affective distractors captured attentional resources more strongly than their neutral counterparts, irrespective of whether they were presented at 6 or 4 Hz rate. Moreover, the emotion-dependent attentional deployment from the foreground task was temporally preceded by sustained sensory facilitation in response to emotional background images. Together, present findings provide evidence for rapid sustained visual facilitation but a rather slow attentional bias in favor of emotional distractors in early visual areas.

Attentional Facilitation of Constituent Features of an Object Does Not Spread Automatically along Object-defining Cortical Boundaries.

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.

It takes two to tango: Suppression of task-irrelevant features requires (spatial) competition.

In a recent electrophysiological study, we reported on global facilitation but local suppression of color stimuli in feature-based attention in human early visual cortex. Subjects attended to one of two centrally located superimposed red/blue random dot kinematograms (RDKs). Task-irrelevant single RDKs in the same colors were presented in the left and right periphery, respectively. Suppression of the to-be-ignored color was only present in the centrally located RDK but not in the one with the same color in the periphery. This result was at odds with the idea of active suppression of task-irrelevant features across the entire visual field. In the present study, we introduced competition in the periphery by superimposing the RDKs at the task-irrelevant location as well. With such competition, we found suppression of the task-irrelevant color in the centrally and peripherally located RDKs. Results clearly demonstrate that suppression of task-irrelevant features at task-irrelevant locations requires (spatial) competitive interactions and is not an inherent neural mechanism in feature-based attention as was found for global facilitation.

Modulation of Somatosensory Alpha Rhythm by Transcranial Alternating Current Stimulation at Mu-Frequency.

Introduction: Transcranial alternating current stimulation (tACS) is emerging as an interventional tool to modulate different functions of the brain, potentially by interacting with intrinsic ongoing neuronal oscillations. Functionally different intrinsic alpha oscillations are found throughout the cortex. Yet it remains unclear whether tACS is capable of specifically modulating the somatosensory mu-rhythm in amplitude. Objectives: We used tACS to modulate mu-alpha oscillations in amplitude. When compared to sham stimulation we expected a modulation of mu-alpha oscillations but not visual alpha oscillations by tACS. Methods: Individual mu-alpha frequencies were determined in 25 participants. Subsequently, blocks of tACS with individual mu-alpha frequency and sham stimulation were applied over primary somatosensory cortex (SI). Electroencephalogram (EEG) was recorded before and after either stimulation or sham. Modulations of mu-alpha and, for control, visual alpha amplitudes were then compared between tACS and sham. Results: Somatosensory mu-alpha oscillations decreased in amplitude after tACS was applied at participants' individual mu-alpha frequency. No changes in amplitude were observed for sham stimulation. Furthermore, visual alpha oscillations were not affected by tACS or sham, respectively. Conclusion: Our results demonstrate the capability of tACS to specifically modulate the targeted somatosensory mu-rhythm when the tACS frequency is tuned to the individual endogenous rhythm and applied over somatosensory areas. Our results are in contrast to previously reported amplitude increases of visual alpha oscillations induced by tACS applied over visual cortex. Our results may point to a specific interaction between our stimulation protocol and the functional architecture of the somatosensory system.

Anodal Transcranial Direct Current Stimulation Does Not Facilitate Dynamic Balance Task Learning in Healthy Old Adults.

Older adults frequently experience a decrease in balance control that leads to increased numbers of falls, injuries and hospitalization. Therefore, evaluating older adults' ability to maintain balance and examining new approaches to counteract age-related decline in balance control is of great importance for fall prevention and healthy aging. Non-invasive brain stimulation techniques such as transcranial direct current stimulation (tDCS) have been shown to beneficially influence motor behavior and motor learning. In the present study, we investigated the influence of tDCS applied over the leg area of the primary motor cortex (M1) on balance task learning of healthy elderly in a dynamic balance task (DBT). In total, 30 older adults were enrolled in a cross-sectional, randomized design including two consecutive DBT training sessions. Only during the first DBT session, either 20 min of anodal tDCS (a-tDCS) or sham tDCS (s-tDCS) were applied and learning improvement was compared between the two groups. Our data showed that both groups successfully learned to perform the DBT on both training sessions. Interestingly, between-group analyses revealed no difference between the a-tDCS and the s-tDCS group regarding their level of task learning. These results indicate that the concurrent application of tDCS over M1 leg area did not elicit DBT learning enhancement in our study cohort. However, a regression analysis revealed that DBT performance can be predicted by the kinematic profile of the movement, a finding that may provide new insights for individualized approaches of treating balance and gait disorders.

Competition for attentional resources between low spatial frequency content of emotional images and a foreground task in early visual cortex.

Low spatial frequency (LSF) image content has been proposed to play a superior functional role in emotional content extraction via the magnocellular pathway biasing attentional resources toward emotional content in visual cortex. We investigated whether emotionally unpleasant complex images that were presented either unfiltered or with LSF content only in the background while subjects performed a foreground task will withdraw more attentional resources from the task compared to unemotional, neutral images (distraction paradigm). We measured steady-state visual evoked potentials (SSVEPs) driven by flickering stimuli of a foreground task. Unfiltered unpleasant images resulted in a significant reduction of SSVEP amplitude compared to neutral images. No statistically significant differences were found with LSF background images. In a behavioral control experiment, we found no significant differences for complexity ratings between unfiltered and LSF pictures. Content identification was possible for unfiltered and LSF picture (correct responses > 74%). An additional EEG study examined typical emotion-related components for complex images presented either as unfiltered, LSF, or high spatial frequency (HSF, as an additional control) filtered, unpleasant, and neutral images. We found a significant main effect of emotional valence in the early posterior negativity. Late positive potential differences were only found for unfiltered and HSF images. Results suggest that, while LSF content is sufficient to allow for content and emotional cue extraction when images were presented alone, LSF content is not salient enough to serve as emotional distractor that withdraws attentional resources from a foreground task in early visual cortex.