The dark side of visual attention

The limited capacity of neural processing restricts the number of objects and locations that can be attended to. Selected events are readily enhanced: the bright side of attention. However, such focal processing comes at a cost, namely, functional blindness for unattended events: the dark side of visual attention. Recent work has advanced our understanding of the neural mechanisms that facilitate visual processing, as well as the neural correlates of unattended, unconscious visual events. Also, new results have revealed how attentional deployment is optimized by non-visual factors such as behavioral set, past experience, and emotional salience.     

Feature-based attentional modulations in the absence of direct visual stimulation

When faced with a crowded visual scene, observers must selectively attend to behaviorally relevant objects to avoid sensory overload. Often this selection process is guided by prior knowledge of a target-defining feature (e.g., the color red when looking for an apple), which enhances the firing rate of visual neurons that are selective for the attended feature. Here, we used functional magnetic resonance imaging and a pattern classification algorithm to predict the attentional state of human observers as they monitored a visual feature (one of two directions of motion). We find that feature-specific attention effects spread across the visual field-even to regions of the scene that do not contain a stimulus. This spread of feature-based attention to empty regions of space may facilitate the perception of behaviorally relevant stimuli by increasing sensitivity to attended features at all locations in the visual field.     

AIM: A network model of attention in auditory cortex

Attentional modulation of cortical networks is critical for the cognitive flexibility required to process complex scenes. Current theoretical frameworks for attention are based almost exclusively on studies in visual cortex, where attentional effects are typically modest and excitatory. In contrast, attentional effects in auditory cortex can be large and suppressive. A theoretical framework for explaining attentional effects in auditory cortex is lacking, preventing a broader understanding of cortical mechanisms underlying attention. Here, we present a cortical network model of attention in primary auditory cortex (A1). A key mechanism in our network is attentional inhibitory modulation (AIM) of cortical inhibitory neurons. In this mechanism, top-down inhibitory neurons disinhibit bottom-up cortical circuits, a prominent circuit motif observed in sensory cortex. Our results reveal that the same underlying mechanisms in the AIM network can explain diverse attentional effects on both spatial and frequency tuning in A1. We find that a dominant effect of disinhibition on cortical tuning is suppressive, consistent with experimental observations. Functionally, the AIM network may play a key role in solving the cocktail party problem. We demonstrate how attention can guide the AIM network to monitor an acoustic scene, select a specific target, or switch to a different target, providing flexible outputs for solving the cocktail party problem.     

视觉注意与神经振荡

人脑每时每刻都要接收大量视觉信息，由于人脑加工信息的能力有限，所以在较大视野内将注意分配给相关信息，同时抑制引起注意分散的不相关信息，对执行目标导向的行为至关重要。这种对视觉信息的选择性和主动性加工以适应当前目标的过程被称作视觉注意（visual attention），且视觉注意可分为自上而下的注意与自下而上的注意两种不同功能。由于来自大脑电信号的神经振荡活动在认知加工中发挥重要作用，已有研究综述了视觉注意与神经振荡（neural oscillation）的密切关系，但并未涉及不同的注意功能与神经振荡的关系。本文系统性调查了不同注意功能与神经振荡的关系，发现额-顶区域的theta频带振荡活动反映了自上而下的认知控制，而后部脑区的theta振荡与自下而上的注意相关。顶-枕区域alpha振荡的偏侧化有助于注意分配，而alpha频带的大规模同步促成了注意对视皮层自上而下的影响。Beta振荡介导了自上而下的信息与自下而上的信息之间的互动，作为信息载体促进了视觉信息处理。Gamma振荡则可能与自上而下和自下而上的注意间整合相关。本文就视觉注意功能与神经振荡关系的研究现状展开综述，旨在揭示不同的神经振荡活动在特定的视觉注意功能中的作用。     

A deficit in covert attention after parietal cortex inactivation in the monkey

Although the parietal cortex has been repeatedly implicated in controlling attention, the nature and importance of this contribution remain unclear. Here we show that inactivating the lateral intraparietal area in monkeys delays the detection of a visual target located in the contralateral visual field. This effect was observed using different visual scene configurations, e.g., with distractors that differ in number or that differ from the target by a conjunction of shape and color or by a single feature. Since eye movements were not allowed during the searching tasks, these results argue for an unambiguous role of the parietal cortex in the top-down control of attentional deployment in space.     

Feature-based attention in the frontal eye field and area V4 during visual search

When we search for a target in a crowded visual scene, we often use the distinguishing features of the target, such as color or shape, to guide our attention and eye movements. To investigate the neural mechanisms of feature-based attention, we simultaneously recorded neural responses in the frontal eye field (FEF) and area V4 while monkeys performed a visual search task. The responses of cells in both areas were modulated by feature attention, independent of spatial attention, and the magnitude of response enhancement was inversely correlated with the number of saccades needed to find the target. However, an analysis of the latency of sensory and attentional influences on responses suggested that V4 provides bottom-up sensory information about stimulus features, whereas the FEF provides a top-down attentional bias toward target features that modulates sensory processing in V4 and that could be used to guide the eyes to a searched-for target.     

Auditory attention--focusing the searchlight on sound

Some fifty years after the first physiological studies of auditory attention, the field is now ripening, with exciting recent insights into the psychophysics, psychology, and neural basis of auditory attention. Current research seeks to unravel the complex interactions of pre-attentive and attentive processing of the acoustic scene, the role of auditory attention in mediating receptive-field plasticity in both auditory spatial and auditory feature processing, the contrasts and parallels between auditory and visual attention pathways and mechanisms, the interplay of bottom-up and top-down attentional mechanisms, the influential role of attention, goals, and expectations in shaping auditory processing, and the orchestration of diverse attentional effects at multiple levels from the cochlea to the cortex.     

Alpha ERD and human visual selective attention

We compared the alpha band EEG depression (event-related desynchnization, ERD) level in two tasks, involving activation of different attentional processes: visual search for a deviant relevant stimulus among many similar ones and visual oddball. Control data for the visual search task consisted of simple viewing of several stimuli being of the same shape as the relevant stimulus in the search trials. Gaze position was verified by eye tracking method. We interpreted alpha band ERD as a correlate of activation of attentional processes. Fixating the target in visual search task caused a significantly larger ERD than fixating the same stimuli in control trials over all leads. We suppose this to be related with task and visual environment complexities. The frontal ERD domination may indicate attentional control over voluntary movements execution (top-down attention). The caudal ERD may be related with updating of visual information as a result of search process (bottom-up attention). Both relevant and irrelevant stimuli in the oddball task also induced alpha band ERD, but it was larger in response to relevant one and reached maximum level over occipital leads. Domination of caudal ERD in oddball task is supposed to indicate bottom-up attention processes.     

Pre-stimulus activity predicts the winner of top-down vs. bottom-up attentional selection

Our ability to process visual information is fundamentally limited. This leads to competition between sensory information that is relevant for top-down goals and sensory information that is perceptually salient, but task-irrelevant. The aim of the present study was to identify, from EEG recordings, pre-stimulus and pre-saccadic neural activity that could predict whether top-down or bottom-up processes would win the competition for attention on a trial-by-trial basis. We employed a visual search paradigm in which a lateralized low contrast target appeared alone, or with a low (i.e., non-salient) or high contrast (i.e., salient) distractor. Trials with a salient distractor were of primary interest due to the strong competition between top-down knowledge and bottom-up attentional capture. Our results demonstrated that 1) in the 1-sec pre-stimulus interval, frontal alpha (8-12 Hz) activity was higher on trials where the salient distractor captured attention and the first saccade (bottom-up win); and 2) there was a transient pre-saccadic increase in posterior-parietal alpha (7-8 Hz) activity on trials where the first saccade went to the target (top-down win). We propose that the high frontal alpha reflects a disengagement of attentional control whereas the transient posterior alpha time-locked to the saccade indicates sensory inhibition of the salient distractor and suppression of bottom-up oculomotor capture.     

Distribution of attention modulates salience signals in early visual cortex

Previous research has shown that the extent to which people spread attention across the visual field plays a crucial role in visual selection and the occurrence of bottom-up driven attentional capture. Consistent with previous findings, we show that when attention was diffusely distributed across the visual field while searching for a shape singleton, an irrelevant salient color singleton captured attention. However, while using the very same displays and task, no capture was observed when observers initially focused their attention at the center of the display. Using event-related fMRI, we examined the modulation of retinotopic activity related to attentional capture in early visual areas. Because the sensory display characteristics were identical in both conditions, we were able to isolate the brain activity associated with exogenous attentional capture. The results show that spreading of attention leads to increased bottom-up exogenous capture and increased activity in visual area V3 but not in V2 and V1.     

Spatiotemporal Dynamics of High-Gamma Activities during a 3-Stimulus Visual Oddball Task

Although many studies have investigated the neural basis of top-down and bottom-up attention, it still requires refinement in both temporal and spatial terms. We used magnetoencephalography to investigate the spatiotemporal dynamics of high-gamma (52–100 Hz) activities during top-down and bottom-up visual attentional processes, aiming to extend the findings from functional magnetic resonance imaging and event-related potential studies. Fourteen participants performed a 3-stimulus visual oddball task, in which both infrequent non-target and target stimuli were presented. We identified high-gamma event-related synchronization in the left middle frontal gyrus, the left intraparietal sulcus, the left thalamus, and the visual areas in different time windows for the target and non-target conditions. We also found elevated imaginary coherence between the left intraparietal sulcus and the right middle frontal gyrus in the high-gamma band from 300 to 400 ms in the target condition, and between the left thalamus and the left middle frontal gyrus in theta band from 150 to 450 ms. In addition, the strength of high-gamma imaginary coherence between the left middle frontal gyrus and left intraparietal sulcus, between the left middle frontal gyrus and the right middle frontal gyrus, and the high-gamma power in the left thalamus predicted inter-subject variation in target detection response time. This source-level electrophysiological evidence enriches our understanding of bi-directional attention processes: stimulus-driven bottom-up attention orientation to a salient, but irrelevant stimulus; and top-down allocation of attentional resources to stimulus evaluation.     

Sensory gain control (amplification) as a mechanism of selective attention: electrophysiological and neuroimaging evidence.

Both physiological and behavioral studies have suggested that stimulus-driven neural activity in the sensory pathways can be modulated in amplitude during selective attention. Recordings of event-related brain potentials indicate that such sensory gain control or amplification processes play an important role in visual-spatial attention. Combined event-related brain potential and neuroimaging experiments provide strong evidence that attentional gain control operates at an early stage of visual processing in extrastriate cortical areas. These data support early selection theories of attention and provide a basis for distinguishing between separate mechanisms of attentional suppression (of unattended inputs) and attentional facilitation (of attended inputs).     

Visual attention: the where,what, how and why of saliency

Attention influences the processing of visual information even in the earliest areas of primate visual cortex. There is converging evidence that the interaction of bottom-up sensory information and top-down attentional influences creates an integrated saliency map, that is, a topographic representation of relative stimulus strength and behavioral relevance across visual space. This map appears to be distributed across areas of the visual cortex, and is closely linked to the oculomotor system that controls eye movements and orients the gaze to locations in the visual scene characterized by a high salience.     

A Neurodynamical Model of Visual Attention: Feedback Enhancement of Spatial Resolution in a Hierarchical System

Human beings have the capacity to recognize objects in natural visual scenes with high efficiency despite the complexity of such scenes, which usually contain multiple objects. One possible mechanism for dealing with this problem is selective attention. Psychophysical evidence strongly suggests that selective attention can enhance the spatial resolution in the input region corresponding to the focus of attention. In this work we adopt a computational neuroscience perspective to analyze the attentional enhancement of spatial resolution in the area containing the objects of interest. We extend and apply the computational model of Deco and Schürmann (2000), which consists of several modules with feedforward and feedback interconnections describing the mutual links between different areas of the visual cortex. Each module analyses the visual input with different spatial resolution and can be thought of as a hierarchical predictor at a given level of resolution. Moreover, each hierarchical predictor has a submodule that consists of a group of neurons performing a biologically based 2D Gabor wavelet transformation at a given resolution level. The attention control decides in which local regions the spatial resolution should be enhanced in a serial fashion. In this sense, the scene is first analyzed at a coarse resolution level, and the focus of attention enhances iteratively the resolution at the location of an object until the object is identified. We propose and simulate new psychophysical experiments where the effect of the attentional enhancement of spatial resolution can be demonstrated by predicting different reaction time profiles in visual search experiments where the target and distractors are defined at different levels of resolution.     

Exploring BOLD Changes during Spatial Attention in Non-Stimulated Visual Cortex

Blood oxygen level-dependent (BOLD) responses were measured in parts of primary visual cortex that represented unstimulated visual field regions at different distances from a stimulated central target location. The composition of the visual scene varied by the presence or absence of additional peripheral distracter stimuli. Bottom-up effects were assessed by comparing peripheral activity during central stimulation vs. no stimulation. Top-down effects were assessed by comparing active vs. passive conditions. In passive conditions subjects simply watched the central letter stimuli and in active conditions they had to report occurrence of pre-defined targets in a rapid serial letter stream. Onset of the central letter stream enhanced activity in V1 representations of the stimulated region. Within representations of the periphery activation decreased and finally turned into deactivation with increasing distance from the stimulated location. This pattern was most pronounced in the active conditions and during the presence of peripheral stimuli. Active search for a target did not lead to additional enhancement at areas representing the attentional focus but to a stronger deactivation in the vicinity. Suppressed neuronal activity was also found in the non distracter condition suggesting a top-down attention driven effect. Our observations suggest that BOLD signal decreases in primary visual cortex are modulated by bottom-up sensory-driven factors such as the presence of distracters in the visual field as well as by top-down attentional processes.     

How does the cerebral cortex work? Learning, attention, and grouping by the laminar circuits of visual cortex

The organization of neocortex into layers is one of its most salient anatomical features. These layers include circuits that form functional columns in cortical maps. A major unsolved problem concerns how bottom-up, top-down, and horizontal interactions are organized within cortical layers to generate adaptive behaviors. This article models how these interactions help visual cortex to realize: (i) the binding process whereby cortex groups distributed data into coherent object representations; (ii) the attentional process whereby cortex selectively processes important events; and (iii) the developmental and learning processes whereby cortex shapes its circuits to match environmental constraints. New computational ideas about feedback systems suggest how neocortex develops and learns in a stable way, and why top-down attention requires converging bottom-up inputs to fully activate cortical cells, whereas perceptual groupings do not.     

Neural circuit models for computations in early visual cortex

Some cortical circuit models study the mechanisms of the transforms from visual inputs to neural responses. They model neural properties such as feature tunings, pattern sensitivities, and how they depend on intracortical connections and contextual inputs. Other cortical circuit models are more concerned with computational goals of the transform from visual inputs to neural responses, or the roles of the neural responses in the visual behavior. The appropriate complexity of a cortical circuit model depends on the question asked. Modeling neural circuits of many interacting hypercolumns is a necessary challenge, which is providing insights to cortical computations, such as visual saliency computation, and linking physiology with global visual cognitive behavior such as bottom-up attentional selection.     

Top-Down but Not Bottom-Up Visual Scanning is Affected in Hereditary Pure Cerebellar Ataxia

The aim of this study was to clarify the nature of visual processing deficits caused by cerebellar disorders. We studied the performance of two types of visual search (top-down visual scanning and bottom-up visual scanning) in 18 patients with pure cerebellar types of spinocerebellar degeneration (SCA6: 11; SCA31: 7). The gaze fixation position was recorded with an eye-tracking device while the subjects performed two visual search tasks in which they looked for a target Landolt figure among distractors. In the serial search task, the target was similar to the distractors and the subject had to search for the target by processing each item with top-down visual scanning. In the pop-out search task, the target and distractor were clearly discernible and the visual salience of the target allowed the subjects to detect it by bottom-up visual scanning. The saliency maps clearly showed that the serial search task required top-down visual attention and the pop-out search task required bottom-up visual attention. In the serial search task, the search time to detect the target was significantly longer in SCA patients than in normal subjects, whereas the search time in the pop-out search task was comparable between the two groups. These findings suggested that SCA patients cannot efficiently scan a target using a top-down attentional process, whereas scanning with a bottom-up attentional process is not affected. In the serial search task, the amplitude of saccades was significantly smaller in SCA patients than in normal subjects. The variability of saccade amplitude (saccadic dysmetria), number of re-fixations, and unstable fixation (nystagmus) were larger in SCA patients than in normal subjects, accounting for a substantial proportion of scattered fixations around the items. Saccadic dysmetria, re-fixation, and nystagmus may play important roles in the impaired top-down visual scanning in SCA, hampering precise visual processing of individual items.     

Increased activity in human visual cortex during directed attention in the absence of visual stimulation

When subjects direct attention to a particular location in a visual scene, responses in the visual cortex to stimuli presented at that location are enhanced, and the suppressive influences of nearby distractors are reduced. What is the top-down signal that modulates the response to an attended versus an unattended stimulus? Here, we demonstrate increased activity related to attention in the absence of visual stimulation in extrastriate cortex when subjects covertly directed attention to a peripheral location expecting the onset of visual stimuli. Frontal and parietal areas showed a stronger signal increase during this expectation than did visual areas. The increased activity in visual cortex in the absence of visual stimulation may reflect a top-down bias of neural signals in favor of the attended location, which derives from a fronto-parietal network.     

A hypothetical role of cortico-basal ganglia-thalamocortical loops in visual processing

The goal of the present work was to define the mechanisms underlying the contribution of sensory and limbic cortico-basal ganglia-thalamocortical loops to visual processing and its attentional modulation. We proposed that visual processing is promoted by dopamine-dependent long-term modifications of synaptic transmission in the basal ganglia that favour a selection of neocortical patterns representing a visual stimulus. This selection is the result of the opposite sign of modulation of strong and weak cortico-basal ganglia inputs and subsequent activity reorganization in each loop. Reorganization leads to disinhibition/inhibition of cortical neurons strongly/weakly excited by stimulus during dopamine release. Recruitment of the thalamo-basal ganglia-collicular pathway is proposed to be necessary for stimulus-evoked dopamine release that underlies bottom-up attentional effects. Visual excitation of the prefrontal cortex and hippocampus (via the thalamus), their cooperation in control of the basal ganglia and dopaminergic cell firing, and simultaneous modulation of activity in diverse cortico-basal ganglia-thalamocortical loops is proposed to underlie top-down attentional effects. It follows from our model that only those components of cortical responses can be modulated by attention, whose onset exceeds the latency of visual responses of dopaminergic cells (50-110 ms). This and other consequences of the model are in accordance with known experimental data.