Distinct Cortical Networks Subserve Spatio-temporal Sampling in Vision through Different Oscillatory Rhythms.

Although visual input arrives continuously, sensory information is segmented into (quasi-)discrete events. Here, we investigated the neural correlates of spatiotemporal binding in humans with magnetoencephalography using two tasks where separate flashes were presented on each trial but were perceived, in a bistable way, as either a single or two separate events. The first task (two-flash fusion) involved judging one versus two flashes, whereas the second task (apparent motion: AM) involved judging coherent motion versus two stationary flashes. Results indicate two different functional networks underlying two unique aspects of temporal binding. In two-flash fusion trials, involving an integration window of ∼50 msec, evoked responses differed as a function of perceptual interpretation by ∼25 msec after stimuli offset. Multivariate decoding of subjective perception based on prestimulus oscillatory phase was significant for alpha-band activity in the right medial temporal (V5/MT) area, with the strength of prestimulus connectivity between early visual areas and V5/MT being predictive of performance. In contrast, the longer integration window (∼130 msec) for AM showed evoked field differences only ∼250 msec after stimuli offset. Phase decoding of the perceptual outcome in AM trials was significant for theta-band activity in the right intraparietal sulcus. Prestimulus theta-band connectivity between V5/MT and intraparietal sulcus best predicted AM perceptual outcome. For both tasks, phase effects found could not be accounted by concomitant variations in power. These results show a strong relationship between specific spatiotemporal binding windows and specific oscillations, linked to the information flow between different areas of the where and when visual pathways.

Accurate localization and coactivation profiles of the frontal eye field and inferior frontal junction: an ALE and MACM fMRI meta-analysis.

The frontal eye field (FEF) and the inferior frontal junction (IFJ) are prefrontal structures involved in mediating multiple aspects of goal-driven behavior. Despite being recognized as prominent nodes of the networks underlying spatial attention and oculomotor control, and working memory and cognitive control, respectively, the limited quantitative evidence on their precise localization has considerably impeded the detailed understanding of their structure and connectivity. In this study, we performed an activation likelihood estimation (ALE) fMRI meta-analysis by selecting studies that employed standard paradigms to accurately infer the localization of these regions in stereotaxic space. For the FEF, we found the highest spatial convergence of activations for prosaccade and antisaccade paradigms at the junction of the precentral sulcus and superior frontal sulcus. For the IFJ, we found consistent activations across oddball/attention, working memory, task-switching and Stroop paradigms at the junction of the inferior precentral sulcus and inferior frontal sulcus. We related these clusters to previous meta-analyses, sulcal/gyral neuroanatomy, and a comprehensive brain parcellation, highlighting important differences compared to their results and taxonomy. Finally, we leveraged the ALE peak coordinates as seeds to perform a meta-analytic connectivity modeling (MACM) analysis, which revealed systematic coactivation patterns spanning the frontal, parietal, and temporal cortices. We decoded the behavioral domains associated with these coactivations, suggesting that these may allow FEF and IFJ to support their specialized roles in flexible behavior. Our study provides the meta-analytic groundwork for investigating the relationship between functional specialization and connectivity of two crucial control structures of the prefrontal cortex.

Functional connectivity fingerprints of the frontal eye field and inferior frontal junction suggest spatial versus nonspatial processing in the prefrontal cortex.

Neuroimaging evidence suggests that the frontal eye field (FEF) and inferior frontal junction (IFJ) govern the encoding of spatial and nonspatial (such as feature- or object-based) representations, respectively, both during visual attention and working memory tasks. However, it is still unclear whether such contrasting functional segregation is also reflected in their underlying functional connectivity patterns. Here, we hypothesized that FEF has predominant functional coupling with spatiotopically organized regions in the dorsal ('where') visual stream whereas IFJ has predominant functional connectivity with the ventral ('what') visual stream. We applied seed-based functional connectivity analyses to temporally high-resolving resting-state magnetoencephalography (MEG) recordings. We parcellated the brain according to the multimodal Glasser atlas and tested, for various frequency bands, whether the spontaneous activity of each parcel in the ventral and dorsal visual pathway has predominant functional connectivity with FEF or IFJ. The results show that FEF has a robust power correlation with the dorsal visual pathway in beta and gamma bands. In contrast, anterior IFJ (IFJa) has a strong power coupling with the ventral visual stream in delta, beta and gamma oscillations. Moreover, while FEF is phase-coupled with the superior parietal lobe in the beta band, IFJa is phase-coupled with the middle and inferior temporal cortex in delta and gamma oscillations. We argue that these intrinsic connectivity fingerprints are congruent with each brain region's function. Therefore, we conclude that FEF and IFJ have dissociable connectivity patterns that fit their respective functional roles in spatial versus nonspatial top-down attention and working memory control.

Decoding Object-Based Auditory Attention from Source-Reconstructed MEG Alpha Oscillations.

How do we attend to relevant auditory information in complex naturalistic scenes? Much research has focused on detecting which information is attended, without regarding underlying top-down control mechanisms. Studies investigating attentional control generally manipulate and cue specific features in simple stimuli. However, in naturalistic scenes it is impossible to dissociate relevant from irrelevant information based on low-level features. Instead, the brain has to parse and select auditory objects of interest. The neural underpinnings of object-based auditory attention remain not well understood. Here we recorded MEG while 15 healthy human subjects (9 female) prepared for the repetition of an auditory object presented in one of two overlapping naturalistic auditory streams. The stream containing the repetition was prospectively cued with 70% validity. Crucially, this task could not be solved by attending low-level features, but only by processing the objects fully. We trained a linear classifier on the cortical distribution of source-reconstructed oscillatory activity to distinguish which auditory stream was attended. We could successfully classify the attended stream from alpha (8-14 Hz) activity in anticipation of repetition onset. Importantly, attention could only be classified from trials in which subjects subsequently detected the repetition, but not from miss trials. Behavioral relevance was further supported by a correlation between classification accuracy and detection performance. Decodability was not sustained throughout stimulus presentation, but peaked shortly before repetition onset, suggesting that attention acted transiently according to temporal expectations. We thus demonstrate anticipatory alpha oscillations to underlie top-down control of object-based auditory attention in complex naturalistic scenes.SIGNIFICANCE STATEMENT In everyday life, we often find ourselves bombarded with auditory information, from which we need to select what is relevant to our current goals. Previous research has highlighted how we attend to specific highly controlled aspects of the auditory input. Although invaluable, it is still unclear how this relates to attentional control in naturalistic auditory scenes. Here we used the high precision of magnetoencephalography in space and time to investigate the brain mechanisms underlying top-down control of object-based attention in ecologically valid sound scenes. We show that rhythmic activity in auditory association cortex at a frequency of ∼10 Hz (alpha waves) controls attention to currently relevant segments within the auditory scene and predicts whether these segments are subsequently detected.

Structure, function and connectivity fingerprints of the frontal eye field versus the inferior frontal junction: A comprehensive comparison.

The human prefrontal cortex contains two prominent areas, the frontal eye field and the inferior frontal junction, that are crucially involved in the orchestrating functions of attention, working memory and cognitive control. Motivated by comparative evidence in non-human primates, we review the human neuroimaging literature, suggesting that the functions of these regions can be clearly dissociated. We found remarkable differences in how these regions relate to sensory domains and visual topography, top-down and bottom-up spatial attention, spatial versus non-spatial (i.e., feature- and object-based) attention and working memory and, finally, the multiple-demand system. Functional magnetic resonance imaging (fMRI) studies using multivariate pattern analysis reveal the selectivity of the frontal eye field and inferior frontal junction to spatial and non-spatial information, respectively. The analysis of functional and effective connectivity provides evidence of the modulation of the activity in downstream visual areas from the frontal eye field and inferior frontal junction and sheds light on their reciprocal influences. We therefore suggest that future studies should aim at disentangling more explicitly the role of these regions in the control of spatial and non-spatial selection. We propose that the analysis of the structural and functional connectivity (i.e., the connectivity fingerprints) of the frontal eye field and inferior frontal junction may be used to further characterize their involvement in a spatial ('where') and a non-spatial ('what') network, respectively, highlighting segregated brain networks that allow biasing visual selection and working memory performance to support goal-driven behaviour.

Author Correction: Cortical source localization of sleep-stage specific oscillatory activity.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Cortical source localization of sleep-stage specific oscillatory activity.

The oscillatory features of non-REM sleep states have been a subject of intense research over many decades. However, a systematic spatial characterization of the spectral features of cortical activity in each sleep state is not available yet. Here, we used magnetoencephalography (MEG) and electroencephalography (EEG) recordings during night sleep. We performed source reconstruction based on the individual subject's anatomical magnetic resonance imaging (MRI) scans and spectral analysis on each non-REM sleep epoch in eight standard frequency bands, spanning the complete spectrum, and computed cortical source reconstructions of the spectral contrasts between each sleep state in comparison to the resting wakefulness. Despite not distinguishing periods of high and low activity within each sleep stage, our results provide new information about relative overall spectral changes in the non-REM sleep stages.

The lateral prefrontal cortex of primates encodes stimulus colors and their behavioral relevance during a match-to-sample task.

The lateral prefrontal cortex of primates (lPFC) plays a central role in complex cognitive behavior, in decision-making as well as in guiding top-down attention. However, how and where in lPFC such behaviorally relevant signals are computed is poorly understood. We analyzed neural recordings from chronic microelectrode arrays implanted in lPFC region 8Av/45 of two rhesus macaques. The animals performed a feature match-to-sample task requiring them to match both motion and color information in a test stimulus. This task allowed to separate the encoding of stimulus motion and color from their current behavioral relevance on a trial-by-trial basis. We found that upcoming motor behavior can be robustly predicted from lPFC activity. In addition, we show that 8Av/45 encodes the color of a visual stimulus, regardless of its behavioral relevance. Most notably, whether a color matches the searched-for color can be decoded independent of a trial's motor outcome and while subjects detect unique feature conjunctions of color and motion. Thus, macaque area 8Av/45 computes, among other task-relevant information, the behavioral relevance of visual color features. Such a signal is most critical for both the selection of responses as well as the deployment of top-down modulatory signals, like feature-based attention.

Alpha Synchrony and the Neurofeedback Control of Spatial Attention.

Decreases in alpha synchronization are correlated with enhanced attention, whereas alpha increases are correlated with inattention. However, correlation is not causality, and synchronization may be a byproduct of attention rather than a cause. To test for a causal role of alpha synchrony in attention, we used MEG neurofeedback to train subjects to manipulate the ratio of alpha power over the left versus right parietal cortex. We found that a comparable alpha asymmetry developed over the visual cortex. The alpha training led to corresponding asymmetrical changes in visually evoked responses to probes presented in the two hemifields during training. Thus, reduced alpha was associated with enhanced sensory processing. Testing after training showed a persistent bias in attention in the expected directions. The results support the proposal that alpha synchrony plays a causal role in modulating attention and visual processing, and alpha training could be used for testing hypotheses about synchrony.

Spatial attention enhances cortical tracking of quasi-rhythmic visual stimuli.

Successfully interpreting and navigating our natural visual environment requires us to track its dynamics constantly. Additionally, we focus our attention on behaviorally relevant stimuli to enhance their neural processing. Little is known, however, about how sustained attention affects the ongoing tracking of stimuli with rich natural temporal dynamics. Here, we used MRI-informed source reconstructions of magnetoencephalography (MEG) data to map to what extent various cortical areas track concurrent continuous quasi-rhythmic visual stimulation. Further, we tested how top-down visuo-spatial attention influences this tracking process. Our bilaterally presented quasi-rhythmic stimuli covered a dynamic range of 4-20 â€‹Hz, subdivided into three distinct bands. As an experimental control, we also included strictly rhythmic stimulation (10 vs 12 â€‹Hz). Using a spectral measure of brain-stimulus coupling, we were able to track the neural processing of left vs. right stimuli independently, even while fluctuating within the same frequency range. The fidelity of neural tracking depended on the stimulation frequencies, decreasing for higher frequency bands. Both attended and non-attended stimuli were tracked beyond early visual cortices, in ventral and dorsal streams depending on the stimulus frequency. In general, tracking improved with the deployment of visuo-spatial attention to the stimulus location. Our results provide new insights into how human visual cortices process concurrent dynamic stimuli and provide a potential mechanism - namely increasing the temporal precision of tracking - for boosting the neural representation of attended input.

Attentional Weighting in the Face Processing Network: A Magnetic Response Image-guided Magnetoencephalography Study Using Multiple Cyclic Entrainments.

We recorded magnetoencephalography using a neural entrainment paradigm with compound face stimuli that allowed for entraining the processing of various parts of a face (eyes, mouth) as well as changes in facial identity. Our magnetic response image-guided magnetoencephalography analyses revealed that different subnodes of the human face processing network were entrained differentially according to their functional specialization. Whereas the occipital face area was most responsive to the rate at which face parts (e.g., the mouth) changed, and face patches in the STS were mostly entrained by rhythmic changes in the eye region, the fusiform face area was the only subregion that was strongly entrained by the rhythmic changes in facial identity. Furthermore, top-down attention to the mouth, eyes, or identity of the face selectively modulated the neural processing in the respective area (i.e., occipital face area, STS, or fusiform face area), resembling behavioral cue validity effects observed in the participants' RT and detection rate data. Our results show the attentional weighting of the visual processing of different aspects and dimensions of a single face object, at various stages of the involved visual processing hierarchy.

Object-based attention in complex, naturalistic auditory streams.

In vision, objects have been described as the 'units' on which non-spatial attention operates in many natural settings. Here, we test the idea of object-based attention in the auditory domain within ecologically valid auditory scenes, composed of two spatially and temporally overlapping sound streams (speech signal vs. environmental soundscapes in Experiment 1 and two speech signals in Experiment 2). Top-down attention was directed to one or the other auditory stream by a non-spatial cue. To test for high-level, object-based attention effects we introduce an auditory repetition detection task in which participants have to detect brief repetitions of auditory objects, ruling out any possible confounds with spatial or feature-based attention. The participants' responses were significantly faster and more accurate in the valid cue condition compared to the invalid cue condition, indicating a robust cue-validity effect of high-level, object-based auditory attention.

Visual Selection of the Future Reach Path in Obstacle Avoidance.

In two EEG experiments, we studied the role of visual attention during the preparation of manual movements around an obstacle. Participants performed rapid hand movements to a goal position avoiding a central obstacle either on the left or right side, depending on the pitch of the acoustical go signal. We used a dot probe paradigm to analyze the deployment of spatial attention in the visual field during the motor preparation. Briefly after the go signal but still before the hand movement actually started, a visual transient was flashed either on the planned pathway of the hand (congruent trials) or on the opposite, movement-irrelevant side (incongruent trials). The P1/N1 components that were evoked by the onset of the dot probe were enhanced in congruent trials where the visual transient was presented on the planned path of the hand. The results indicate that, during movement preparation, attention is allocated selectively to the planned trajectory the hand is going to take around the obstacle.

Electrical stimulation of macaque lateral prefrontal cortex modulates oculomotor behavior indicative of a disruption of top-down attention.

The lateral prefrontal cortex (lPFC) of primates is hypothesized to be heavily involved in decision-making and selective visual attention. Recent neurophysiological evidence suggests that information necessary for an orchestration of those high-level cognitive factors are indeed represented in the lPFC. However, we know little about the specific contribution of sub-networks within lPFC to the deployment of top-down influences that can be measured in extrastriate visual cortex. Here, we systematically applied electrical stimulations to areas 8Av and 45 of two macaque monkeys performing a concurrent goal-directed saccade task. Despite using currents well above saccadic thresholds of the directly adjacent Frontal Eye Fields (FEF), saccades were only rarely evoked by the stimulation. Instead, two types of behavioral effects were observed: Stimulations of caudal sites in 8Av (close to FEF) shortened or prolonged saccadic reaction times, depending on the task-instructed saccade, while rostral stimulations of 8Av/45 seem to affect the relative attentional weighting of saccade targets as well as saccadic reaction times. These results illuminate important differences in the causal involvement of different sub-networks within the lPFC and are most compatible with a stimulation-induced biasing of stimulus processing that accelerates the detection of saccade targets presented ipsilateral to stimulation through a disruption of contralaterally deployed top-down attention.

Distinct distributed patterns of neural activity are associated with two languages in the bilingual brain.

A large body of previous neuroimaging studies suggests that multiple languages are processed and organized in a single neuroanatomical system in the bilingual brain, although differential activation may be seen in some studies because of different proficiency levels and/or age of acquisition of the two languages. However, one important possibility is that the two languages may involve interleaved but functionally independent neural populations within a given cortical region, and thus, distinct patterns of neural computations may be pivotal for the processing of the two languages. Using functional magnetic resonance imaging (fMRI) and multivariate pattern analyses, we tested this possibility in Chinese-English bilinguals when they performed an implicit reading task. We found a broad network of regions wherein the two languages evoked different patterns of activity, with only partially overlapping patterns of voxels in a given region. These regions, including the middle occipital cortices, fusiform gyri, and lateral temporal, temporoparietal, and prefrontal cortices, are associated with multiple aspects of language processing. The results suggest the functional independence of neural computations underlying the representations of different languages in bilinguals.

Neural mechanisms of object-based attention.

How we attend to objects and their features that cannot be separated by location is not understood. We presented two temporally and spatially overlapping streams of objects, faces versus houses, and used magnetoencephalography and functional magnetic resonance imaging to separate neuronal responses to attended and unattended objects. Attention to faces versus houses enhanced the sensory responses in the fusiform face area (FFA) and parahippocampal place area (PPA), respectively. The increases in sensory responses were accompanied by induced gamma synchrony between the inferior frontal junction, IFJ, and either FFA or PPA, depending on which object was attended. The IFJ appeared to be the driver of the synchrony, as gamma phases were advanced by 20 ms in IFJ compared to FFA or PPA. Thus, the IFJ may direct the flow of visual processing during object-based attention, at least in part through coupled oscillations with specialized areas such as FFA and PPA.

Chunking movements into sequence: the visual pre-selection of subsequent goals.

The chunking of individual movements into sequences has been studied extensively from a motor point of view. Here we approach the planning of sequential behavior from a perceptual perspective investigating the sensorimotor transformations that accompany visually guided sequential behavior. We show that visual attention pre-selects subsequent goals only if two movements are planned to be carried out in rapid succession and therefore are integrated into one common action. This causes visual attention to select both intended goal locations in advance. In contrast, in more slowly executed motor sequences, the single movements are programmed one-by-one and subsequent movement goals are only later visually prepared ('just in time'). The visual selection of a subsequent goal location crucially depends on the speed of the planned sequence: the longer the inter-reach delay, the less visual attention is deployed to the subsequent goal initially.

Attentional landscapes in reaching and grasping.

It is well established that during the preparation and execution of goal-directed movements, perceptual processing is biased towards the goal. Most of the previous work on the relation between action and attention has focused on rather simple movements, such as single saccades or manual reaches towards a single target. Here we review recent behavioural and neurophysiological studies on manual actions that require to consider more than a single spatial location in the planning of the response, such as movement sequences, grasping, and movements around obstacles. The studies provide compelling evidence that the preparation of these actions establishes multiple foci of attention which reflect the spatial-temporal requirements of the future action. The findings help clarify how perceptual processing is bound by action preparation and also offer new perspectives for motor control research.

Time perception as a workload measure in simulated car driving.

In experimental studies using flight simulations subjects' duration estimates have shown to be an effective indicator of cognitive task demands. In this study we wanted to find out whether subjective time perception could serve as a measure of cognitive workload during simulated car driving. Participants drove on a round course of a driving simulator consisting of three different environments with different levels of task demands. Drivers were required to perform a time-production task while driving the vehicle. Electrodermal activity and subjective ratings of mental workload (SWAT) were recorded simultaneously. The length of produced intervals increased significantly in more complex driving situations, as did electrodermal activity and subjective ratings of mental workload. Thus, time production is a valid indicator of cognitive involvement in simulated driving and could become a valid method to measure the current mental workload of car drivers in various traffic situations.

Attentional selection of multiple goal positions before rapid hand movement sequences: an event-related potential study.

A dot-probe paradigm was used to provide physiological evidence for the parallel selection of multiple movement goals before rapid hand movement sequences. Participants executed a sequence of manual pointing movements to two out of three possible goal positions. During movement preparation, a task-irrelevant visual transient (a dot probe) was flashed either at one of both movement goals, or at the third, movement-irrelevant location. The results revealed that the N1 component induced by the presentation of the dot was enhanced if the dot was flashed at one of the movement goals, indicating that both target positions were attended before the initialization of the movement sequence. A second experiment showed that movement-irrelevant locations between the movement goals were not attended, suggesting that attention splits into spatially distinct foci.