The neural basis of attentional selection in goal-directed memory retrieval.

Goal-directed memory reactivation involves retrieving the most relevant information for the current behavioral goal. Previous research has linked this process to activations in the fronto-parietal network, but the underlying neurocognitive mechanism remains poorly understood. The current electroencephalogram (EEG) study explores attentional selection as a possible mechanism supporting goal-directed retrieval. We designed a long-term memory experiment containing three phases. First, participants learned associations between objects and two screen locations. In a following phase, we changed the relevance of some locations (selective cue condition) to simulate goal-directed retrieval. We also introduced a control condition, in which the original associations remained unchanged (neutral cue condition). Behavior performance measured during the final retrieval phase revealed faster and more confident responses in the selective vs. neutral condition. At the EEG level, we found significant differences in decoding accuracy, with above-chance effects in the selective cue condition but not in the neutral cue condition. Additionally, we observed a stronger posterior contralateral negativity and lateralized alpha power in the selective cue condition. Overall, these results suggest that attentional selection enhances task-relevant information accessibility, emphasizing its role in goal-directed memory retrieval.

Search flavor labels in beverages: An electrophysiological investigation of color-flavor congruency and association strength in visual search.

Individuals are apt to link various characteristics of an object or event through different sensory experiences. We conducted two electrophysiological experiments to investigate the effects of color-flavor congruency and association strength on visual search efficiency and the in-depth cognitive mechanisms underlying multisensory processes. Participants were prompted with a flavor label and asked to identify the primed flavor from four beverage bottle images. Experiment 1 focused on color-flavor congruency and noted faster searches for congruent targets than incongruent ones. EEG data exhibited smaller N2, larger P3 and LPC, and increased parietal-occipital midline (POM) alpha power for incongruent targets than congruent ones. Experiment 2 manipulated color-flavor association strength within each flavor. Behavioral findings showed that searches for targets with weak association strength took longer than those with strong association strength. Moreover, time-frequency analysis displayed that the former evoked greater frontal midline (FM) theta power and higher alpha power than the latter. Altogether, our research indicated that (1) color expectations based on prior experience can automatically guide people's attentional selection, (2) the color-flavor congruency and association strength impact the visual search efficiency via distinct pathways, and (3) theta and alpha activities make a pivotal role in unraveling multisensory information processing. These findings shed some light on the intricate cognitive processes involved in crossmodal visual search and the underlying neurocognitive dynamics.

Division and spreading of attention across color.

Biological systems must allocate limited perceptual resources to relevant elements in their environment. This often requires simultaneous selection of multiple elements from the same feature dimension (e.g. color). To establish the determinants of divided attentional selection of color, we conducted an experiment that used multicolored displays with four overlapping random dot kinematograms that differed only in hue. We manipulated (i) requirement to focus attention to a single color or divide it between two colors; (ii) distances of distractor hues from target hues in a perceptual color space. We conducted a behavioral and an electroencephalographic experiment, in which each color was tagged by a specific flicker frequency and driving its own steady-state visual evoked potential. Behavioral and neural indices of attention showed several major consistencies. Concurrent selection halved the neural signature of target enhancement observed for single targets, consistent with an approximately equal division of limited resources between two hue-selective foci. Distractors interfered with behavioral performance in a context-dependent fashion but their effects were asymmetric, indicating that perceptual distance did not adequately capture attentional distance. These asymmetries point towards an important role of higher-level mechanisms such as categorization and grouping-by-color in determining the efficiency of attentional allocation in complex, multicolored scenes.

Peripheral vision is mainly for looking rather than seeing.

Vision includes looking and seeing. Looking, mainly via gaze shifts, selects a fraction of visual input information for passage through the brain's information bottleneck. The selected input is placed within the attentional spotlight, typically in the central visual field. Seeing decodes, i.e., recognizes and discriminates, the selected inputs. Hence, peripheral vision should be mainly devoted to looking, in particular, deciding where to shift the gaze. Looking is often guided exogenously by a saliency map created by the primary visual cortex (V1), and can be effective with no seeing and limited awareness. In seeing, peripheral vision not only suffers from poor spatial resolution, but is also subject to crowding and is more vulnerable to illusions by misleading, ambiguous, and impoverished visual inputs. Central vision, mainly for seeing, enjoys the top-down feedback that aids seeing in light of the bottleneck which is hypothesized to starts from V1 to higher areas. This feedback queries for additional information from lower visual cortical areas such as V1 for ongoing recognition. Peripheral vision is deficient in this feedback according to the Central-peripheral Dichotomy (CPD) theory. The saccades engendered by peripheral vision allows looking to combine with seeing to give human observers the impression of seeing the whole scene clearly despite inattentional blindness.

Top-down control of exogenous attentional selection is mediated by beta coherence in prefrontal cortex.

Salience-driven exogenous and goal-driven endogenous attentional selection are two distinct forms of attention that guide selection of task-irrelevant and task-relevant targets in primates. Top-down attentional control mechanisms enable selection of the task-relevant target by limiting the influence of sensory information. Although the lateral prefrontal cortex (LPFC) is known to mediate top-down control, the neuronal mechanisms of top-down control of attentional selection are poorly understood. Here, we trained two rhesus monkeys on a two-target, free-choice luminance-reward selection task. We demonstrate that visual-movement (VM) neurons and nonvisual neurons or movement neurons encode exogenous and endogenous selection. We then show that coherent beta activity selectively modulates mechanisms of exogenous selection specifically during conflict and consequently may support top-down control. These results reveal the VM-neuron-specific network mechanisms of attentional selection and suggest a functional role for beta-frequency coherent neural dynamics in the modulation of sensory communication channels for the top-down control of attentional selection.

Good-enough attentional guidance.

Theories of attention posit that attentional guidance operates on information held in a target template within memory. The template is often thought to contain veridical target features, akin to a photograph, and to guide attention to objects that match the exact target features. However, recent evidence suggests that attentional guidance is highly flexible and often guided by non-veridical features, a subset of features, or only associated features. We integrate these findings and propose that attentional guidance maximizes search efficiency based on a 'good-enough' principle to rapidly localize candidate target objects. Candidates are then serially interrogated to make target-match decisions using more precise information. We suggest that good-enough guidance optimizes the speed-accuracy-effort trade-offs inherent in each stage of visual search.

Attention and platypuses.

This perspective piece discusses a set of attentional phenomena that are not easily accommodated within current theories of attentional selection. We call these phenomena attentional platypuses, as they allude to an observation that within biological taxonomies the platypus does not fit into either mammal or bird categories. Similarly, attentional phenomena that do not fit neatly within current attentional models suggest that current models are in need of a revision. We list a few instances of the "attentional platypuses" and then offer a new approach, that we term dynamically weighted prioritization, stipulating that multiple factors impinge onto the attentional priority map, each with a corresponding weight. The interaction between factors and their corresponding weights determines the current state of the priority map which subsequently constrains/guides attentional allocation. We propose that this new approach should be considered as a supplement to existing models of attention, especially those that emphasize categorical organizations. This article is categorized under: Psychology > Attention Psychology > Perception and Psychophysics Neuroscience > Cognition.

Information-based multivariate decoding reveals imprecise neural encoding in children with attention deficit hyperactivity disorder during visual selective attention.

Attention deficit hyperactivity disorder (ADHD) is a common neurodevelopmental disorder in school-age children. Attentional orientation is a potential clinical diagnostic marker to aid in the early diagnosis of ADHD. However, the underlying pathophysiological substrates of impaired attentional orienting in childhood ADHD remain unclear. Electroencephalography (EEG) was measured in 135 school-age children (70 with childhood ADHD and 65 matched typically developing children) to directly investigate target localization during spatial selective attention through univariate ERP analysis and information-based multivariate pattern machine learning analysis. Compared with children with typical development, a smaller N2pc was found in the ADHD group through univariate ERP analysis. Children with ADHD showed a lower parieto-occipital multivariate decoding accuracy approximately 240-340 ms after visual search onset, which predicts a slower reaction time and larger standard deviation of reaction time. Furthermore, a significant correlation was found between N2pc and decoding accuracy in typically developing children but not in children with ADHD. These observations reveal that impaired attentional orienting in ADHD may be due to inefficient neural encoding responses. By using a personalized information-based multivariate machine learning approach, we have advanced the understanding of cognitive deficits in neurodevelopmental disorders. Our study provides potential research directions for the early diagnosis and optimization of personalized intervention in children with ADHD.

Attention in Schizophrenia.

Attention is clearly a core area of cognitive dysfunction in schizophrenia, but the concept of "attention" is complex and multifaceted. This chapter focuses on three different aspects of attentional function that are of particular interest in schizophrenia. First, we discuss the evidence that schizophrenia involves a reduction in global alertness, leading to an inward focusing of attention and a neglect of external stimuli and tasks. Second, we discuss the control of attention, the set of processes that allow general goals to be translated into shifts of attention toward task-relevant information. When a goal is adequately represented, people with schizophrenia often show no deficit in using the goal to direct attention in the visual modality unless challenged by stimuli that strongly activate the magnocellular processing pathway. Finally, we discuss the implementation of selection, the processes that boost relevant information and suppress distractors once attention has been directed to a given source of information. Although early evidence indicated an impairment in selection, more recent evidence indicating that people with schizophrenia actually focus their attention more narrowly and more intensely that healthy individuals (hyperfocusing). However, this hyperfocused attention may be directed toward goal-irrelevant information, creating the appearance of impaired attentional filtering.

What to expect where and when: how statistical learning drives visual selection.

While the visual environment contains massive amounts of information, we should not and cannot pay attention to all events. Instead, we need to direct attention to those events that have proven to be important in the past and suppress those that were distracting and irrelevant. Experiences molded through a learning process enable us to extract and adapt to the statistical regularities in the world. While previous studies have shown that visual statistical learning (VSL) is critical for representing higher order units of perception, here we review the role of VSL in attentional selection. Evidence suggests that through VSL, attentional priority settings are optimally adjusted to regularities in the environment, without intention and without conscious awareness.

Narrative Comprehension Guides Eye Movements in the Absence of Motion.

Viewers' attentional selection while looking at scenes is affected by both top-down and bottom-up factors. However, when watching film, viewers typically attend to the movie similarly irrespective of top-down factors-a phenomenon we call the tyranny of film. A key difference between still pictures and film is that film contains motion, which is a strong attractor of attention and highly predictive of gaze during film viewing. The goal of the present study was to test if the tyranny of film is driven by motion. To do this, we created a slideshow presentation of the opening scene of Touch of Evil. Context condition participants watched the full slideshow. No-context condition participants did not see the opening portion of the scene, which showed someone placing a time bomb into the trunk of a car. In prior research, we showed that despite producing very different understandings of the clip, this manipulation did not affect viewers' attention (i.e., the tyranny of film), as both context and no-context participants were equally likely to fixate on the car with the bomb when the scene was presented as a film. The current study found that when the scene was shown as a slideshow, the context manipulation produced differences in attentional selection (i.e., it attenuated attentional synchrony). We discuss these results in the context of the Scene Perception and Event Comprehension Theory, which specifies the relationship between event comprehension and attentional selection in the context of visual narratives.

Aging impairs primary task resumption and attentional control processes following interruptions.

Attentional selection of working memory content is impaired after an interruption. Here we investigate the neural correlates underlying attentional selection within working memory. We focus especially on how older and younger adults differ in attentional selection processes during primary task resumption. Participants performed a working memory task, while being frequently interrupted with either a cognitively low- or high-demanding arithmetic task. Afterwards, a retrospective cue (retro-cue) indicated the working memory content required for later report. The detrimental effect of the interruption was evident in both age groups, but while younger adults were more strongly affected by a high- than by a low-demanding interruption, the performance deficit appeared independently of the cognitive requirements of the interruption task in older adults. A similar pattern was found regarding frontal-posterior connectivity in the theta frequency range, suggesting that aging decreases the ability to selectively maintain relevant information within working memory. The power of mid-frontal theta oscillations (~4-9 Hz) featured a comparable effect of interruptions in both age groups. However, posterior alpha/beta power (~8-30 Hz) following the retro-cue was more diminished by a preceding interruption in older adults. These results suggest an age-related deficit in the attentional selection and maintenance of primary task information following an interruption that appeared independent from the cognitive requirements of the interrupting task.

Statistical learning in visual search reflects distractor rarity, not only attentional suppression.

In visual search tasks, salient distractors may capture attention involuntarily, but interference can be reduced when the salient distractor appears more frequently on one out of several possible positions. The reduction was attributed to attentional suppression of the high-probability position. However, all previous studies on this topic compared performance on the high-probability position to the remaining positions, which had a low probability of containing the distractor. Therefore, it is not clear whether the difference resulted from reduced interference on the high-probability position or from increased interference on the low-probability positions. To decide between these alternatives, we compared high-probability and low-probability with equal-probability positions. Consistent with attentional suppression, interference was reduced on the high-probability position compared with equal-probability positions. However, there was also an increase in interference on low-probability positions compared with equal-probability positions. The increase is in line with previous reports of boosted interference when distractors are rare. Our results show that the experimental design used in previous research is insufficient to separate effects of attentional suppression and those of distractor rarity.

The diachronic account of attentional selectivity.

Many models of attention assume that attentional selection takes place at a specific moment in time that demarcates the critical transition from pre-attentive to attentive processing of sensory input. We argue that this intuitively appealing standard account of attentional selectivity is not only inaccurate, but has led to substantial conceptual confusion. As an alternative, we offer a 'diachronic' framework that describes attentional selectivity as a process that unfolds over time. Key to this view is the concept of attentional episodes, brief periods of intense attentional amplification of sensory representations that regulate access to working memory and response-related processes. We describe how attentional episodes are linked to earlier attentional mechanisms and to recurrent processing at the neural level. We review studies that establish the existence of attentional episodes, delineate the factors that determine if and when they are triggered, and discuss the costs associated with processing multiple events within a single episode. Finally, we argue that this framework offers new solutions to old problems in attention research that have never been resolved. It can provide a unified and conceptually coherent account of the network of cognitive and neural processes that produce the goal-directed selectivity in perceptual processing that is commonly referred to as 'attention'.

Preparatory attention incorporates contextual expectations.

Humans are remarkably proficient at finding objects within complex visual scenes. According to current theories of attention,1-3 visual processing of an object of interest is favored through the preparatory activation of object-specific representations in visual cortex.4-15 One key problem that is inherent to real-world visual search but is not accounted for by current theories is that a given object will produce a dramatically different retinal image depending on its location, which is unknown in advance. For instance, the color of the retinal image depends on the illumination on the object, its shape depends on the viewpoint, and (most critically) its size can vary by several orders of magnitude, depending on the distance to the observer. In order to benefit search, preparatory activity thus needs to incorporate contextual expectations. In the current study, we measured fMRI blood-oxygen-level-dependent (BOLD) activity in human observers while they prepared to search for objects at different distances in indoor-scene photographs. First, we established that observers instantiated preparatory object representations: activity patterns in object-selective cortex evoked during search preparation (while no objects were presented) resembled activity patterns evoked by viewing those objects in isolation. Second, we demonstrated that these preparatory object representations were systematically modulated by expectations derived from scene context: activity patterns reflected the predicted retinal image of the object at each distance (i.e., distant search evoking smaller object representations and nearby search evoking larger object representations). These findings reconcile current theories of attentional selection with the challenges of real-world vision.

History Modulates Early Sensory Processing of Salient Distractors.

To find important objects, we must focus on our goals, ignore distractions, and take our changing environment into account. This is formalized in models of visual search whereby goal-driven, stimulus-driven, and history-driven factors are integrated into a priority map that guides attention. Stimulus history robustly influences where attention is allocated even when the physical stimulus is the same: when a salient distractor is repeated over time, it captures attention less effectively. A key open question is how we come to ignore salient distractors when they are repeated. Goal-driven accounts propose that we use an active, expectation-driven mechanism to attenuate the distractor signal (e.g., predictive coding), whereas stimulus-driven accounts propose that the distractor signal is attenuated because of passive changes to neural activity and inter-item competition (e.g., adaptation). To test these competing accounts, we measured item-specific fMRI responses in human visual cortex during a visual search task where trial history was manipulated (colors unpredictably switched or were repeated). Consistent with a stimulus-driven account of history-based distractor suppression, we found that repeated singleton distractors were suppressed starting in V1, and distractor suppression did not increase in later visual areas. In contrast, we observed signatures of goal-driven target enhancement that were absent in V1, increased across visual areas, and were not modulated by stimulus history. Our data suggest that stimulus history does not alter goal-driven expectations, but rather modulates canonically stimulus-driven sensory responses to contribute to a temporally integrated representation of priority.SIGNIFICANCE STATEMENT Visual search refers to our ability to find what we are looking for in a cluttered visual world (e.g., finding your keys). To perform visual search, we must integrate information about our goals (e.g., "find the red keychain"), the environment (e.g., salient items capture your attention), and changes to the environment (i.e., stimulus history). Although stimulus history impacts behavior, the neural mechanisms that mediate history-driven effects remain debated. Here, we leveraged fMRI and multivariate analysis techniques to measure history-driven changes to the neural representation of items during visual search. We found that stimulus history influenced the representation of a salient "pop-out" distractor starting in V1, suggesting that stimulus history operates via modulations of early sensory processing rather than goal-driven expectations.

Across-trial spatial suppression in visual search.

In order to focus on objects of interest, humans must be able to avoid distraction by salient stimuli that are not relevant to the task at hand. Many recent studies have shown that through statistical learning we are able to suppress the location that is most likely to contain a salient distractor. Here we demonstrate a remarkable flexibility in attentional suppression. Participants had to search for a shape singleton while a color distractor singleton was present. Unbeknown to the participant, the color distractor was presented according to a consistent pattern across trials. Our findings show that participants learn this distractor sequence as they proactively suppressed the anticipated location of the distractor on the next trial. Critically, none of the participants were aware of these hidden sequences. We conclude that the spatial priority map is highly flexible, operating at a subconscious level preparing the attentional system for what will happen next.

Neurochemistry of Visual Attention.

Visual attention is the cognitive process that mediates the selection of important information from the environment. This selection is usually controlled by bottom-up and top-down attentional biasing. Since for most humans vision is the dominant sense, visual attention is critically important for higher-order cognitive functions and related deficits are a core symptom of many neuropsychiatric and neurological disorders. Here, we summarize the importance and relative contributions of different neuromodulators and neurotransmitters to the neural mechanisms of top-down and bottom-up attentional control. We will not only review the roles of widely accepted neuromodulators, such as acetylcholine, dopamine and noradrenaline, but also the contributions of other modulatory substances. In doing so, we hope to shed some light on the current understanding of the role of neurochemistry in shaping neuron properties contributing to the allocation of attention in the visual field.

Differing Time Courses of Reward-Related Attentional Processing: An EEG Source-Space Analysis.

Since our environment typically contains more information than can be processed at any one time due to the limited capacity of our visual system, we are bound to differentiate between relevant and irrelevant information. This process, termed attentional selection, is usually categorized into bottom-up and top-down processes. However, recent research suggests reward might also be an important factor in guiding attention. Monetary reward can bias attentional selection in favor of task-relevant targets and reduce the efficiency of visual search when a reward-associated, but task-irrelevant distractor is present. This study is the first to investigate reward-related target and distractor processing in an additional singleton task using neurophysiological measures and source space analysis. Based on previous studies, we hypothesized that source space analysis would find enhanced neural activity in regions of the value-based attention network, such as the visual cortex and the anterior cingulate. Additionally, we went further and explored the time courses of the underlying attentional mechanisms. Our neurophysiological results showed that rewarding distractors led to a stronger attentional capture. In line with this, we found that reward-associated distractors (compared with reward-associated targets) enhanced activation in frontal regions, indicating the involvement of top-down control processes. As hypothesized, source space analysis demonstrated that reward-related targets and reward-related distractors elicited activation in regions of the value-based attention network. However, these activations showed time-dependent differences, indicating that the neural mechanisms underlying reward biasing might be different for task-relevant and task-irrelevant stimuli.

Representational dynamics preceding conscious access.

Our senses are continuously bombarded with more information than our brain can process up to the level of awareness. The present study aimed to enhance understanding on how attentional selection shapes conscious access under conditions of rapidly changing input. Using an attention task, EEG, and multivariate decoding of individual target- and distractor-defining features, we specifically examined dynamic changes in the representation of targets and distractors as a function of conscious access and the task-relevance (target or distractor) of the preceding item in the RSVP stream. At the behavioral level, replicating previous work and suggestive of a flexible gating mechanism, we found a significant impairment in conscious access to targets (T2) that were preceded by a target (T1) followed by one or two distractors (i.e., the attentional blink), but striking facilitation of conscious access to targets shown directly after another target (i.e., lag-1 sparing and blink reversal). At the neural level, conscious access to T2 was associated with enhanced early- and late-stage T1 representations and enhanced late-stage D1 representations, and interestingly, could be predicted based on the pattern of EEG activation well before T1 was presented. Yet, across task conditions, we did not find convincing evidence for the notion that conscious access is affected by rapid top-down selection-related modulations of the strength of early sensory representations induced by the preceding visual event. These results cannot easily be explained by existing accounts of how attentional selection shapes conscious access under rapidly changing input conditions, and have important implications for theories of the attentional blink and consciousness more generally.