Top-down control of the left visual field bias in cued visual spatial attention.

A left visual field (LVF) bias in perceptual judgments, response speed, and discrimination accuracy has been reported in humans. Cognitive factors, such as visual spatial attention, are known to modulate or even eliminate this bias. We investigated this problem by recording pupillometry together with functional magnetic resonance imaging (fMRI) in a cued visual spatial attention task. We observed that (i) the pupil was significantly more dilated following attend-right than attend-left cues, (ii) the task performance (e.g. reaction time [RT]) did not differ between attend-left and attend-right trials, and (iii) the difference in cue-related pupil dilation between attend-left and attend-right trials was inversely related to the corresponding difference in RT. Neuroscientically, correlating the difference in cue-related pupil dilation with the corresponding cue-related fMRI difference yielded activations primarily in the right hemisphere, including the right intraparietal sulcus and the right ventrolateral prefrontal cortex. These results suggest that (i) there is an asymmetry in visual spatial attention control, with the rightward attention control being more effortful than the leftward attention control, (ii) this asymmetry underlies the reduction or the elimination of the LVF bias, and (iii) the components of the attentional control networks in the right hemisphere are likely part of the neural substrate of the observed asymmetry in attentional control.

Time Courses of Attended and Ignored Object Representations.

Selective attention prioritizes information that is relevant to behavioral goals. Previous studies have shown that attended visual information is processed and represented more efficiently, but distracting visual information is not fully suppressed, and may also continue to be represented in the brain. In natural vision, to-be-attended and to-be-ignored objects may be present simultaneously in the scene. Understanding precisely how each is represented in the visual system, and how these neural representations evolve over time, remains a key goal in cognitive neuroscience. In this study, we recorded EEG while participants performed a cued object-based attention task that involved attending to target objects and ignoring simultaneously presented and spatially overlapping distractor objects. We performed support vector machine classification on the stimulus-evoked EEG data to separately track the temporal dynamics of target and distractor representations. We found that (1) both target and distractor objects were decodable during the early phase of object processing (∼100 msec to ∼200 msec after target onset), and (2) the representations of both objects were sustained over time, remaining decodable above chance until ∼1000-msec latency. However, (3) the distractor object information faded significantly beginning after about 300-msec latency. These findings provide information about the fate of attended and ignored visual information in complex scene perception.

Role of Inferior Frontal Junction (IFJ) in the Control of Feature versus Spatial Attention.

Feature-based visual attention refers to preferential selection and processing of visual stimuli based on their nonspatial attributes, such as color or shape. Recent studies have highlighted the inferior frontal junction (IFJ) as a control region for feature but not spatial attention. However, the extent to which IFJ contributes to spatial versus feature attention control remains a topic of debate. We investigated in humans of both sexes the role of IFJ in the control of feature versus spatial attention in a cued visual spatial (attend-left or attend-right) and feature (attend-red or attend-green) attention task using fMRI. Analyzing cue-related fMRI using both univariate activation and multivoxel pattern analysis, we found the following results in IFJ. First, in line with some prior studies, the univariate activations were not different between feature and spatial attentional control. Second, in contrast, the multivoxel pattern analysis decoding accuracy was above chance level for feature attention (attend-red vs attend-green) but not for spatial attention (attend-left vs attend-right). Third, while the decoding accuracy for feature attention was above chance level during attentional control in the cue-to-target interval, it was not during target processing. Fourth, the right IFJ and visual cortex (V4) were observed to be functionally connected during feature but not during spatial attention control, and this functional connectivity was positively associated with subsequent attentional selection of targets in V4, as well as with behavioral performance. These results support a model in which IFJ plays a crucial role in top-down control of visual feature but not visual spatial attention.SIGNIFICANCE STATEMENT Past work has shown that the inferior frontal junction (IFJ), a prefrontal structure, is activated by both attention-to-feature (e.g., color) and attention-to-location, but the precise role of IFJ in the control of feature- versus spatial-attention is debated. We investigated this issue in a cued visual spatial (attend-left or attend-right) and feature (attend-red or attend-green) attention task using fMRI, multivoxel pattern analysis, and functional connectivity methods. The results show that (1) attend-red versus attend-green can be decoded in single-trial cue-evoked BOLD activity in IFJ but not attend-left versus attend-right and (2) only right IFJ modulates V4 to enhance task performance. This study sheds light on the function and hemispheric specialization of IFJ in the control of visual attention.

Neural Mechanisms of Attentional Control for Objects: Decoding EEG Alpha When Anticipating Faces, Scenes,and Tools.

Attentional selection mechanisms in visual cortex involve changes in oscillatory activity in the EEG alpha band (8-12 Hz), with decreased alpha indicating focal cortical enhancement and increased alpha indicating suppression. This has been observed for spatial selective attention and attention to stimulus features such as color versus motion. We investigated whether attention to objects involves similar alpha-mediated changes in focal cortical excitability. In experiment 1, 20 volunteers (8 males; 12 females) were cued (80% predictive) on a trial-by-trial basis to different objects (faces, scenes, or tools). Support vector machine decoding of alpha power patterns revealed that late (>500 ms latency) in the cue-to-target foreperiod, only EEG alpha differed with the to-be-attended object category. In experiment 2, to eliminate the possibility that decoding of the physical features of cues led to our results, 25 participants (9 males; 16 females) performed a similar task where cues were nonpredictive of the object category. Alpha decoding was now only significant in the early (<200 ms) foreperiod. In experiment 3, to eliminate the possibility that task set differences between the different object categories led to our experiment 1 results, 12 participants (5 males; 7 females) performed a predictive cuing task where the discrimination task for different objects was identical across object categories. The results replicated experiment 1. Together, these findings support the hypothesis that the neural mechanisms of visual selective attention involve focal cortical changes in alpha power not only for simple spatial and feature attention, but also for high-level object attention in humans.SIGNIFICANCE STATEMENT Attention is the cognitive function that enables relevant information to be selected from sensory inputs so it can be processed in the support of goal-directed behavior. Visual attention is widely studied, yet the neural mechanisms underlying the selection of visual information remain unclear. Oscillatory EEG activity in the alpha range (8-12 Hz) of neural populations receptive to target visual stimuli may be part of the mechanism, because alpha is thought to reflect focal neural excitability. Here, we show that alpha-band activity, as measured by scalp EEG from human participants, varies with the specific category of object selected by attention. This finding supports the hypothesis that alpha-band activity is a fundamental component of the neural mechanisms of attention.

The Microstructure of Attentional Control in the Dorsal Attention Network.

The top-down control of attention involves command signals arising chiefly in the dorsal attention network (DAN) in frontal and parietal cortex and propagating to sensory cortex to enable the selective processing of incoming stimuli based on their behavioral relevance. Consistent with this view, the DAN is active during preparatory (anticipatory) attention for relevant events and objects, which, in vision, may be defined by different stimulus attributes including their spatial location, color, motion, or form. How this network is organized to support different forms of preparatory attention to different stimulus attributes remains unclear. We propose that, within the DAN, there exist functional microstructures (patterns of activity) specific for controlling attention based on the specific information to be attended. To test this, we contrasted preparatory attention to stimulus location (spatial attention) and to stimulus color (feature attention), and used multivoxel pattern analysis to characterize the corresponding patterns of activity within the DAN. We observed different multivoxel patterns of BOLD activation within the DAN for the control of spatial attention (attending left vs. right) and feature attention (attending red vs. green). These patterns of activity for spatial and feature attentional control showed limited overlap with each other within the DAN. Our findings thus support a model in which the DAN has different functional microstructures for distinctive forms of top-down control of visual attention.

Theta Oscillations Index Frontal Decision-Making and Mediate Reciprocal Frontal-Parietal Interactions in Willed Attention.

Attention can be attracted reflexively by sensory signals, biased by learning or reward, or focused voluntarily based on momentary goals. When voluntary attention is focused by purely internal decision processes (will), rather than instructions via external cues, we call this "willed attention." In prior work, we reported ERP and fMRI correlates of willed spatial attention in trial-by-trial cuing tasks. Here we further investigated the oscillatory mechanisms of willed attention by contrasting the event-related EEG spectrogram between instructional and choice cues. Two experiments were conducted at 2 different sites using the same visuospatial attention paradigm. Consistent between the 2 experiments, we found increases in frontal theta power (starting at ~500 ms post cue) for willed attention relative to instructed attention. This frontal theta increase was accompanied by increased frontal-parietal theta-band coherence and bidirectional Granger causality. Additionally, the onset of attention-related posterior alpha power lateralization was delayed in willed attention relative to instructed attention, and the amount of delay was related to the timing of frontal theta increase. These results, replicated across 2 experiments, suggest that theta oscillations are the neuronal signals indexing decision-making in the frontal cortex, and mediating reciprocal communications between the frontal executive and parietal attentional control regions during willed attention.

Attention enhances synaptic efficacy and the signal-to-noise ratio in neural circuits.

Attention is a critical component of perception. However, the mechanisms by which attention modulates neuronal communication to guide behaviour are poorly understood. To elucidate the synaptic mechanisms of attention, we developed a sensitive assay of attentional modulation of neuronal communication. In alert monkeys performing a visual spatial attention task, we probed thalamocortical communication by electrically stimulating neurons in the lateral geniculate nucleus of the thalamus while simultaneously recording shock-evoked responses from monosynaptically connected neurons in primary visual cortex. We found that attention enhances neuronal communication by increasing the efficacy of presynaptic input in driving postsynaptic responses, by increasing synchronous responses among ensembles of postsynaptic neurons receiving independent input, and by decreasing redundant signals between postsynaptic neurons receiving common input. The results demonstrate that attention finely tunes neuronal communication at the synaptic level by selectively altering synaptic weights, enabling enhanced detection of salient events in the noisy sensory environment.

Deciding where to attend: Large-scale network mechanisms underlying attention and intention revealed by graph-theoretic analysis.

The neural mechanisms by which intentions are transformed into actions remain poorly understood. We investigated the network mechanisms underlying spontaneous voluntary decisions about where to focus visual-spatial attention (willed attention). Graph-theoretic analysis of two independent datasets revealed that regions activated during willed attention form a set of functionally-distinct networks corresponding to the frontoparietal network, the cingulo-opercular network, and the dorsal attention network. Contrasting willed attention with instructed attention (where attention is directed by external cues), we observed that the dorsal anterior cingulate cortex was allied with the dorsal attention network in instructed attention, but shifted connectivity during willed attention to interact with the cingulo-opercular network, which then mediated communications between the frontoparietal network and the dorsal attention network. Behaviorally, greater connectivity in network hubs, including the dorsolateral prefrontal cortex, the dorsal anterior cingulate cortex, and the inferior parietal lobule, was associated with faster reaction times. These results, shown to be consistent across the two independent datasets, uncover the dynamic organization of functionally-distinct networks engaged to support intentional acts.

Top-down Modulation of Neural Activity in Anticipatory Visual Attention: Control Mechanisms Revealed by Simultaneous EEG-fMRI.

In covert visual attention, frontoparietal attention control areas are thought to issue signals to selectively bias sensory neurons to facilitate behaviorally relevant information and suppress distraction. We investigated the relationship between activity in attention control areas and attention-related modulation of posterior alpha activity using simultaneous electroencephalography (EEG) and functional magnetic resonance imaging in humans during cued visual-spatial attention. Correlating single-trial EEG alpha power with blood-oxygen-level dependent (BOLD) activity, we found that BOLD in the intraparietal sulcus (IPS) and left middle frontal gyrus was inversely correlated with occipital alpha power. Importantly, in IPS, inverse correlations were stronger for alpha within the hemisphere contralateral to the attended hemifield, implicating the IPS in the enhancement of task-relevant sensory areas. Positive BOLD-alpha correlations were observed in sensorimotor cortices and the default mode network, suggesting a mechanism of active suppression over task-irrelevant areas. The magnitude of cue-induced alpha lateralization was positively correlated with BOLD in dorsal anterior cingulate cortex and dorsolateral prefrontal cortex, implicating a role of executive control in attention. These results show that IPS and frontal executive areas are the main sources of biasing influences on task-relevant visual cortex, whereas task-irrelevant default mode network and sensorimotor cortex are inhibited during visual attention.

Electrophysiological Evidence for Impaired Control of Motor Output in Schizophrenia.

Previous research has demonstrated pervasive deficits in response-related processing in people with schizophrenia (PSZ). The present study used behavioral measures and event-related potentials (ERPs) to test the hypothesis that schizophrenia involves specific impairment in the ability to exert control over response-related processing. Twenty-two PSZ and 22 matched control participants completed a choice response task in counterbalanced testing sessions that emphasized only accuracy (the unspeeded condition) or emphasized speed and accuracy equally (the speeded condition). Control participants successfully modulated behavioral and ERP indices of response-related processing under speed pressure, as evidenced by faster and less variable reaction times (RTs) and an earlier onset and increased amplitude lateralized readiness potential (LRP). By contrast, PSZ were unable to improve RT speed or variability or to modulate the LRP under speed pressure, despite showing a decrease in accuracy. Notably, response-related deficits in PSZ emerged only in the speeded condition; behavioral and ERP measures did not differ between groups in the unspeeded condition. Together, these results indicate that impairment in the ability to exert control over response-related processing may underlie response-related deficits in schizophrenia.

ERP Evidence for Implicit Priming of Top-Down Control of Attention.

Proportion congruency effects are the observation that the magnitude of the Stroop effect increases as the proportion of congruent trials in a block increases. Contemporary work shows that proportion effects can be specific to a particular context. For example, in a Simon task in which items appearing above fixation are mostly congruent and items appearing below fixation are mostly incongruent, the Simon effect is larger for the items appearing at the top. There is disagreement as to whether these context-specific effects result from simple associative learning or, instead, a type of conflict-mediated associative learning. Here, we address this question in an ERP study using a Simon task in which the proportion congruency effect was context-specific, manipulating the proportion of congruent trials based on location (upper vs. lower visual field). We found significant behavioral proportion congruency effects that varied with the specific contexts. In addition, we observed that the N2 response of the ERPs to the stimuli was larger in amplitude for the high congruent (high conflict) versus low congruent (low conflict) conditions/contexts. Because the N2 is known to be greater in amplitude also for trials where conflict is high and is believed to be an electrical signal related to conflict detection in the medial frontal cortex, this supports the idea that conflict-mediated associative learning is involved in the proportion congruency effect.

Simultaneous recordings from the primary visual cortex and lateral geniculate nucleus reveal rhythmic interactions and a cortical source for γ-band oscillations.

Oscillatory synchronization of neuronal activity has been proposed as a mechanism to modulate effective connectivity between interacting neuronal populations. In the visual system, oscillations in the gamma-frequency range (30-100 Hz) are thought to subserve corticocortical communication. To test whether a similar mechanism might influence subcortical-cortical communication, we recorded local field potential activity from retinotopically aligned regions in the lateral geniculate nucleus (LGN) and primary visual cortex (V1) of alert macaque monkeys viewing stimuli known to produce strong cortical gamma-band oscillations. As predicted, we found robust gamma-band power in V1. In contrast, visual stimulation did not evoke gamma-band activity in the LGN. Interestingly, an analysis of oscillatory phase synchronization of LGN and V1 activity identified synchronization in the alpha (8-14 Hz) and beta (15-30 Hz) frequency bands. Further analysis of directed connectivity revealed that alpha-band interactions mediated corticogeniculate feedback processing, whereas beta-band interactions mediated geniculocortical feedforward processing. These results demonstrate that although the LGN and V1 display functional interactions in the lower frequency bands, gamma-band activity in the alert monkey is largely an emergent property of cortex.

Sensitivity to Referential Ambiguity in Discourse: The Role of Attention, Working Memory, and Verbal Ability.

The establishment of reference is essential to language comprehension. The goal of this study was to examine listeners' sensitivity to referential ambiguity as a function of individual variation in attention, working memory capacity, and verbal ability. Participants listened to stories in which two entities were introduced that were either very similar (e.g., two oaks) or less similar (e.g., one oak and one elm). The manipulation rendered an anaphor in a subsequent sentence (e.g., oak) ambiguous or unambiguous. EEG was recorded as listeners comprehended the story, after which participants completed tasks to assess working memory, verbal ability, and the ability to use context in task performance. Power in the alpha and theta frequency bands when listeners received critical information about the discourse entities (e.g., oaks) was used to index attention and the involvement of the working memory system in processing the entities. These measures were then used to predict an ERP component that is sensitive to referential ambiguity, the Nref, which was recorded when listeners received the anaphor. Nref amplitude at the anaphor was predicted by alpha power during the earlier critical sentence: Individuals with increased alpha power in ambiguous compared with unambiguous stories were less sensitive to the anaphor's ambiguity. Verbal ability was also predictive of greater sensitivity to referential ambiguity. Finally, increased theta power in the ambiguous compared with unambiguous condition was associated with higher working-memory span. These results highlight the role of attention and working memory in referential processing during listening comprehension.

Normal aging selectively diminishes alpha lateralization in visual spatial attention.

EEG studies of cue-induced visual alpha power (8-13 Hz) lateralization have been conducted on young adults without examining differences that may develop as a consequence of normal aging. Here, we examined age-related differences in spatial attention by comparing healthy older and younger adults. Our key finding is that cue-induced alpha power lateralization was observed in younger, but not older adults, even though both groups exhibited classic event-related potential signatures of spatial orienting. Specifically, both younger and older adults showed significant early directing-attention negativity (EDAN), anterior directing-attention negativity (ADAN), late directing-attention positivity (LDAP) and contingent negative variation (CNV). Furthermore, target-evoked sensory components were enhanced for attended relative to unattended targets in both younger and older groups. This pattern of results suggests that although older adults can successfully allocate spatial attention, they do so without the lateralization of alpha power that is commonly observed in younger adults. Taken together, our findings demonstrate that younger and older adults might engage different neural mechanisms for attentional orienting, and that alpha power lateralization during visual spatial attention is a phenomenon that diminishes during normal aging.

The neural correlates of volitional attention: A combined fMRI and ERP study.

Studies of visual-spatial attention typically use instructional cues to direct attention to a relevant location, but in everyday vision, attention is often focused volitionally, in the absence of external signals. Although investigations of cued attention comprise hundreds of behavioral and physiological studies, remarkably few studies of voluntary attention have addressed the challenging question of how spatial attention is initiated and controlled in the absence of external instructions, which we refer to as willed attention. To explore this question, we employed a trial-by-trial spatial attention task using electroencephalography and functional magnetic resonance imaging (fMRI). The fMRI results reveal a unique network of brain regions for willed attention that includes the anterior cingulate cortex, left middle frontal gyrus (MFG), and the left and right anterior insula (AI). We also observed two event-related potentials (ERPs) associated with willed attention; one with a frontal distribution occurring 250-350 ms postdecision cue onset (EWAC: Early Willed Attention Component), and another occurring between 400 and 800 ms postdecision-cue onset (WAC: Willed Attention Component). In addition, each ERP component uniquely correlated across subjects with different willed attention-specific sites of BOLD activation. The EWAC was correlated with the willed attention-specific left AI and left MFG activations and the later WAC was correlated only with left AI. These results offer a comprehensive and novel view of the electrophysiological and anatomical profile of willed attention and further illustrate the relationship between scalp-recorded ERPs and the BOLD response.

Spared and impaired spoken discourse processing in schizophrenia: effects of local and global language context.

Individuals with schizophrenia are impaired in a broad range of cognitive functions, including impairments in the controlled maintenance of context-relevant information. In this study, we used ERPs in human subjects to examine whether impairments in the controlled maintenance of spoken discourse context in schizophrenia lead to overreliance on local associations among the meanings of individual words. Healthy controls (n = 22) and patients (n = 22) listened to short stories in which we manipulated global discourse congruence and local priming. The target word in the last sentence of each story was globally congruent or incongruent and locally associated or unassociated. ERP local association effects did not significantly differ between control participants and schizophrenia patients. However, in contrast to controls, patients only showed effects of discourse congruence when targets were primed by a word in the local context. When patients had to use discourse context in the absence of local priming, they showed impaired brain responses to the target. Our findings indicate that schizophrenia patients are impaired during discourse comprehension when demands on controlled maintenance of context are high. We further found that ERP measures of increased reliance on local priming predicted reduced social functioning, suggesting that alterations in the neural mechanisms underlying discourse comprehension have functional consequences in the illness.

Spontaneous neural fluctuations predict decisions to attend.

Ongoing variability in neural signaling is an intrinsic property of the brain. Often this variability is considered to be noise and ignored. However, an alternative view is that this variability is fundamental to perception and cognition and may be particularly important in decision-making. Here, we show that a momentary measure of occipital alpha-band power (8-13 Hz) predicts choices about where human participants will focus spatial attention on a trial-by-trial basis. This finding provides evidence for a mechanistic account of decision-making by demonstrating that ongoing neural activity biases voluntary decisions about where to attend within a given moment.

The role of the pulvinar in distractor processing and visual search.

The pulvinar nuclei of the thalamus are hypothesized to coordinate attentional selection in the visual cortex. Different models have, however, been proposed for the precise role of the pulvinar in attention. One proposal is that the pulvinar mediates shifts of spatial attention; a different proposal is that it serves the filtering of distractor information. At present, the relation between these possible operations and their relative importance in the pulvinar remains unresolved. We address this issue by contrasting these proposals in two fMRI experiments. We used a visual search paradigm that permitted us to dissociate neural activity reflecting shifts of attention from activity underlying distractor filtering. We find that distractor filtering, but not the operation of shifting attention, is associated with strong activity enhancements in dorsal and ventral regions of the pulvinar as well as in early visual cortex areas including the primary visual cortex. Our observations indicate that distractor filtering is the preponderant attentional operation subserved by the pulvinar, presumably mediated by a modulation of processing in visual areas where spatial resolution is sufficiently high to separate target from distractor input.

Optimizing cognitive neuroscience experiments for separating event- related fMRI BOLD responses in non-randomized alternating designs.

Functional magnetic resonance imaging (fMRI) has revolutionized human brain research. But there exists a fundamental mismatch between the rapid time course of neural events and the sluggish nature of the fMRI blood oxygen level-dependent (BOLD) signal, which presents special challenges for cognitive neuroscience research. This limitation in the temporal resolution of fMRI puts constraints on the information about brain function that can be obtained with fMRI and also presents methodological challenges. Most notably, when using fMRI to measure neural events occurring closely in time, the BOLD signals may temporally overlap one another. This overlap problem may be exacerbated in complex experimental paradigms (stimuli and tasks) that are designed to manipulate and isolate specific cognitive-neural processes involved in perception, cognition, and action. Optimization strategies to deconvolve overlapping BOLD signals have proven effective in providing separate estimates of BOLD signals from temporally overlapping brain activity, but there remains reduced efficacy of such approaches in many cases. For example, when stimulus events necessarily follow a non-random order, like in trial-by-trial cued attention or working memory paradigms. Our goal is to provide guidance to improve the efficiency with which the underlying responses evoked by one event type can be detected, estimated, and distinguished from other events in designs common in cognitive neuroscience research. We pursue this goal using simulations that model the nonlinear and transient properties of fMRI signals, and which use more realistic models of noise. Our simulations manipulated: (i) Inter-Stimulus-Interval (ISI), (ii) proportion of so-called null events, and (iii) nonlinearities in the BOLD signal due to both cognitive and design parameters. We offer a theoretical framework along with a python toolbox called deconvolve to provide guidance on the optimal design parameters that will be of particular utility when using non-random, alternating event sequences in experimental designs. In addition, though, we also highlight the challenges and limitations in simultaneously optimizing both detection and estimation efficiency of BOLD signals in these common, but complex, cognitive neuroscience designs.

Top-Down Biasing of Visual Cortical Activity Encodes Attended Information and Facilitates Behavioral Performance in Visual Spatial Attention.

Top-down attention plays a vital role in selecting relevant stimuli and suppressing distracting information. During top-down visual-spatial attention, control signals from the dorsal attention network modulate the baseline neuronal activity in the visual cortex in favor of task-relevant stimuli. While several studies have demonstrated that baseline shift during anticipatory attention occurs in multiple visual areas, such effects have not been systematically investigated across the visual hierarchy, especially when different attention conditions are matched for stimulus and task factors. In this fMRI study, we investigated anticipatory attention signals using univariate and multivariate (MVPA) analysis in multiple visual cortical areas. First, the univariate analysis yielded significant activation differences in higher-order visual areas, with the effect weaker in early visual areas. Second, however, in contrast, MVPA decoding was significant in predicting attention conditions in all visual areas and IPS, with lower-order visual areas (e.g., V1) having greater decoding accuracy than higher-order visual areas (e.g., LO1). Third, the strength of decoding accuracy predicted the behavioral performance in the discrimination task. All the results were highly replicable and consistent across two datasets with same experimental paradigms but recorded at two research sites, and two experimental conditions where the direction of spatial attention was driven either by external instructions (cue-instructed attention) or from internal decisions (free-choice attention). Our results provide clear evidence, not available in past univariate investigations, that top-down attentional control signals selectively bias neuronal processing throughout the visual hierarchy, and that this biasing is correlated with the task performance.