Parallel attentional facilitation of features and objects in early visual cortex.
ABSTRACT: Selective attention can enhance the processing of attended features across the entire visual field. Attention also spreads within objects, enhancing all internal locations and task-irrelevant features of selected objects. Here, we examine the extent to which attentional enhancement of a feature spreads across attended and unattended objects. Two fully overlapping counter-rotating bicolored surfaces of light and dark random dots were presented on a gray background of intermediate luminance. This stimulus creates a percept of two separate semitransparent surfaces and allows the measurement of feature- and object-based selections while controlling spatial attention. On each trial, human participants attended to a subset of dots defined by feature (luminance polarity) and object (surface) in order to detect brief episodes of radial motion while ignoring any events in the unattended groups of dots. Attentional selection was assessed by means of steady-state visual evoked potentials (SSVEPs) and behavioral measures. SSVEP amplitudes recorded at medial occipital electrode sites were modulated both by surface-based and luminance polarity-based selection in a manner consistent with independent multiplicative enhancement of attentional effects in different dimensions in early visual cortex. This finding supports the view that feature-based attention spreads across object boundaries, at least at an early stage of processing. However, SSVEPs elicited at more lateral electrode sites showed a hierarchical pattern of selection, potentially reflecting the binding of surface-defining features with luminance features to enable surface-based attention.
Project description:Finding and recognizing objects is a fundamental task of vision. Objects can be defined by several "cues" (color, luminance, texture, etc.), and humans can integrate sensory cues to improve detection and recognition [1-3]. Cortical mechanisms fuse information from multiple cues , and shape-selective neural mechanisms can display cue invariance by responding to a given shape independent of the visual cue defining it [5-8]. Selective attention, in contrast, improves recognition by isolating a subset of the visual information . Humans can select single features (red or vertical) within a perceptual dimension (color or orientation), giving faster and more accurate responses to items having the attended feature [10, 11]. Attention elevates neural responses and sharpens neural tuning to the attended feature, as shown by studies in psychophysics and modeling [11, 12], imaging [13-16], and single-cell and neural population recordings [17, 18]. Besides single features, attention can select whole objects [19-21]. Objects are among the suggested "units" of attention because attention to a single feature of an object causes the selection of all of its features [19-21]. Here, we pit integration against attentional selection in object recognition. We find, first, that humans can integrate information near optimally from several perceptual dimensions (color, texture, luminance) to improve recognition. They cannot, however, isolate a single dimension even when the other dimensions provide task-irrelevant, potentially conflicting information. For object recognition, it appears that there is mandatory integration of information from multiple dimensions of visual experience. The advantage afforded by this integration, however, comes at the expense of attentional selection.
Project description:Contrasting theories of visual attention have emphasized selection by spatial location, individual features, and whole objects. We used functional magnetic resonance imaging to ask whether and how attention to one feature of an object spreads to other features of the same object. Subjects viewed two spatially superimposed surfaces of random dots that were segregated by distinct color-motion conjunctions. The color and direction of motion of each surface changed smoothly and in a cyclical fashion. Subjects were required to track one feature (e.g., color) of one of the two surfaces and detect brief moments when the attended feature diverged from its smooth trajectory. To tease apart the effect of attention to individual features on the hemodynamic response, we used a frequency-tagging scheme. In this scheme, the stimulus features (color and direction of motion) are modulated periodically at distinct frequencies so that the contribution of each feature to the hemodynamics can be inferred from the harmonic response at the corresponding frequency. We found that attention to one feature (e.g., color) of one surface increased the response modulation not only to the attended feature but also to the other feature (e.g., motion) of the same surface. This attentional modulation was evident in multiple visual areas and was present as early as V1. The spread of attention to the behaviorally irrelevant features of a surface suggests that attention may automatically select all features of a single object. Thus object-based attention may be supported by an enhancement of feature-specific sensory signals in the visual cortex.
Project description:Humans can flexibly select locations, features, or objects in a visual scene for prioritized processing. Although it is relatively straightforward to manipulate location- and feature-based attention, it is difficult to isolate object-based selection. Because objects are always composed of features, studies of object-based selection can often be interpreted as the selection of a combination of locations and features. Here we examined the neural representation of attentional priority in a paradigm that isolated object-based selection. Participants viewed two superimposed gratings that continuously changed their color, orientation, and spatial frequency, such that the gratings traversed the same exact feature values within a trial. Participants were cued at the beginning of each trial to attend to one or the other grating to detect a brief luminance increment, while their brain activity was measured with fMRI. Using multi-voxel pattern analysis, we were able to decode the attended grating in a set of frontoparietal areas, including anterior intraparietal sulcus (IPS), frontal eye field (FEF), and inferior frontal junction (IFJ). Thus, a perceptually varying object can be represented by patterned neural activity in these frontoparietal areas. We suggest that these areas can encode attentional priority for abstract, high-level objects independent of their locations and features.
Project description:Recognizing objects in cluttered scenes requires attentional mechanisms to filter out distracting information. Previous studies have found several physiological correlates of attention in visual cortex, including larger responses for attended objects. However, it has been unclear whether these attention-related changes have a large impact on information about objects at the neural population level. To address this question, we trained monkeys to covertly deploy their visual attention from a central fixation point to one of three objects displayed in the periphery, and we decoded information about the identity and position of the objects from populations of ? 200 neurons from the inferior temporal cortex using a pattern classifier. The results show that before attention was deployed, information about the identity and position of each object was greatly reduced relative to when these objects were shown in isolation. However, when a monkey attended to an object, the pattern of neural activity, represented as a vector with dimensionality equal to the size of the neural population, was restored toward the vector representing the isolated object. Despite this nearly exclusive representation of the attended object, an increase in the salience of nonattended objects caused "bottom-up" mechanisms to override these "top-down" attentional enhancements. The method described here can be used to assess which attention-related physiological changes are directly related to object recognition, and should be helpful in assessing the role of additional physiological changes in the future.
Project description:In a crowded visual scene, attention must be distributed efficiently and flexibly over time and space to accommodate different contexts. It is well established that selective attention enhances the corresponding neural responses, presumably implying that attention would persistently dwell on the task-relevant item. Meanwhile, recent studies, mostly in divided attentional contexts, suggest that attention does not remain stationary but samples objects alternately over time, suggesting a rhythmic view of attention. However, it remains unknown whether the dynamic mechanism essentially mediates attentional processes at a general level. Importantly, there is also a complete lack of direct neural evidence reflecting whether and how the brain rhythmically samples multiple visual objects during stimulus processing. To address these issues, in this study, we employed electroencephalography (EEG) and a temporal response function (TRF) approach, which can dissociate responses that exclusively represent a single object from the overall neuronal activity, to examine the spatiotemporal characteristics of attention in various attentional contexts. First, attention, which is characterized by inhibitory alpha-band (approximately 10 Hz) activity in TRFs, switches between attended and unattended objects every approximately 200 ms, suggesting a sequential sampling even when attention is required to mostly stay on the attended object. Second, the attentional spatiotemporal pattern is modulated by the task context, such that alpha-mediated switching becomes increasingly prominent as the task requires a more uniform distribution of attention. Finally, the switching pattern correlates with attentional behavioral performance. Our work provides direct neural evidence supporting a generally central role of temporal organization mechanism in attention, such that multiple objects are sequentially sorted according to their priority in attentional contexts. The results suggest that selective attention, in addition to the classically posited attentional "focus," involves a dynamic mechanism for monitoring all objects outside of the focus. Our findings also suggest that attention implements a space (object)-to-time transformation by acting as a series of concatenating attentional chunks that operate on 1 object at a time.
Project description:Faced with an overwhelming amount of sensory information, we are able to prioritize the processing of select spatial locations and visual features. The neuronal mechanisms underlying such spatial and feature-based selection have been studied in considerable detail. More recent work shows that attention can also be allocated to objects, even spatially superimposed objects composed of dynamically changing features that must be integrated to create a coherent object representation. Much less is known about the mechanisms underlying such object-based selection. Our goal was to investigate behavioral and neuronal responses when attention was directed to one of two objects, specifically one of two superimposed transparent surfaces, in a task designed to preclude space-based and feature-based selection. We used functional magnetic resonance imaging (fMRI) to measure changes in blood oxygen level-dependent (BOLD) signals when attention was deployed to one or the other surface. We found that visual areas V1, V2, V3, V3A, and MT+ showed enhanced BOLD responses to translations of an attended relative to an unattended surface. These results reveal that visual areas as early as V1 can be modulated by attending to objects, even objects defined by dynamically changing elements. This provides definitive evidence in humans that early visual areas are involved in a seemingly high-order process. Furthermore, our results suggest that these early visual areas may participate in object-specific feature "binding," a process that seemingly must occur for an object or a surface to be the unit of attentional selection.
Project description:Spatial attention must adjust around an object of interest in a manner that reflects the object's size on the retina as well as the proximity of distracting objects, a process often guided by nonspatial features. This study used ERPs to investigate how quickly the size of this type of "attentional window" can adjust around a fixated target object defined by its color and whether this variety of attention influences the feedforward flow of subsequent information through the visual system. The task involved attending either to a circular region at fixation or to a surrounding annulus region, depending on which region contained an attended color. The region containing the attended color varied randomly from trial to trial, so the spatial distribution of attention had to be adjusted on each trial. We measured the initial sensory ERP response elicited by an irrelevant probe stimulus that appeared in one of the two regions at different times after task display onset. This allowed us to measure the amount of time required to adjust spatial attention on the basis of the location of the task-relevant feature. We found that the probe-elicited sensory response was larger when the probe occurred within the region of the attended dots, and this effect required a delay of approximately 175 msec between the onset of the task display and the onset of the probe. Thus, the window of attention is rapidly adjusted around the point of fixation in a manner that reflects the spatial extent of a task-relevant stimulus, leading to changes in the feedforward flow of subsequent information through the visual system.
Project description:Visual object perception requires integration of multiple features; spatial attention is thought to be critical to this binding. But attention is rarely static-how does dynamic attention impact object integrity? Here, we manipulated covert spatial attention and had participants (total N = 48) reproduce multiple properties (color, orientation, location) of a target item. Object-feature binding was assessed by applying probabilistic models to the joint distribution of feature errors: Feature reports for the same object could be correlated (and thus bound together) or independent. We found that splitting attention across multiple locations degrades object integrity, whereas rapid shifts of spatial attention maintain bound objects. Moreover, we document a novel attentional phenomenon, wherein participants exhibit unintentional fluctuations- lapses of spatial attention-yet nevertheless preserve object integrity at the wrong location. These findings emphasize the importance of a single focus of spatial attention for object-feature binding, even when that focus is dynamically moving across the visual field.
Project description:Theories of object-based attention often make two assumptions: that attentional resources are facilitatory, and that they spread automatically within grouped objects. Consistent with this, ignored visual stimuli can be easier to process, or more distracting, when perceptually grouped with an attended target stimulus. But in past studies, the ignored stimuli often shared potentially relevant features or locations with the target. In this fMRI study, we measured the effects of attention and grouping on Blood Oxygenation Level Dependent (BOLD) responses in the human brain to entirely task-irrelevant events. Two checkerboards were displayed each in opposite hemifields, while participants responded to check-size changes in one pre-cued hemifield, which varied between blocks. Grouping (or segmentation) between hemifields was manipulated between blocks, using common (vs. distinct) motion cues. Task-irrelevant transient events were introduced by randomly changing the color of either checkerboard, attended or ignored, at unpredictable intervals. The above assumptions predict heightened BOLD signals for irrelevant events in attended vs. ignored hemifields for ungrouped contexts, but less such attentional modulation under grouping, due to automatic spreading of facilitation across hemifields. We found the opposite pattern, in primary visual cortex. For ungrouped stimuli, BOLD signals associated with task-irrelevant changes were lower, not higher, in the attended vs. ignored hemifield; furthermore, attentional modulation was not reduced but actually inverted under grouping, with higher signals for events in the attended vs. ignored hemifield. These results challenge two popular assumptions underlying object-based attention. We consider a broader biased-competition framework: task-irrelevant stimuli are suppressed according to how strongly they compete with task-relevant stimuli, with intensified competition when the irrelevant features or locations comprise the same object.
Project description:How attentional modulation on brain activities determines behavioral performance has been one of the most important issues in cognitive neuroscience. This issue has been addressed by comparing the temporal relationship between attentional modulations on neural activities and behavior. Our previous study measured the time course of attention with amplitude and phase coherence of steady-state visual evoked potential (SSVEP) and found that the modulation latency of phase coherence rather than that of amplitude was consistent with the latency of behavioral performance. In this study, as a complementary report, we compared the time course of visual attention shift measured by event-related potentials (ERPs) with that by target detection task. We developed a novel technique to compare ERPs with behavioral results and analyzed the EEG data in our previous study. Two sets of flickering stimulus at different frequencies were presented in the left and right visual hemifields, and a target or distracter pattern was presented randomly at various moments after an attention-cue presentation. The observers were asked to detect targets on the attended stimulus after the cue. We found that two ERP components, P300 and N2pc, were elicited by the target presented at the attended location. Time-course analyses revealed that attentional modulation of the P300 and N2pc amplitudes increased gradually until reaching a maximum and lasted at least 1.5 s after the cue onset, which is similar to the temporal dynamics of behavioral performance. However, attentional modulation of these ERP components started later than that of behavioral performance. Rather, the time course of attentional modulation of behavioral performance was more closely associated with that of the concurrently recorded SSVEPs analyzed. These results suggest that neural activities reflected not by either the P300 or N2pc, but by the SSVEPs, are the source of attentional modulation of behavioral performance.