Humans can attend to different objects independent of their spatial locations. While selecting an object has been shown to modulate object processing in high-level visual areas in occipitotemporal cortex, where/how behavioral importance (i.e., priority) for objects is represented is unknown. Here we examined the patterns of distributed neural activity during an object-based selection task. We measured brain activity with functional magnetic resonance imaging (fMRI), while participants viewed two superimposed, dynamic objects (left- and right-pointing triangles) and were cued to attend to one of the triangle objects. Enhanced fMRI response was observed for the attention conditions compared to a neutral condition, but no significant difference was found in overall response amplitude between two attention conditions. By using multi-voxel pattern classification (MVPC), however, we were able to distinguish the neural patterns associated with attention to different objects in early visual cortex (V1 to hMT+) and lateral occipital complex (LOC). Furthermore, distinct multi-voxel patterns were also observed in frontal and parietal areas. Our results demonstrate that object-based attention has a wide-spread modulation effect along the visual hierarchy and suggest that object-specific priority information is represented by patterned neural activity in the dorsal frontoparietal network.
Although considerable research has exa mined the storage limits of visual short-term memory (VSTM), little is known ab out the initial formation (i.e., the consolidation) of VSTM representations. A few previous studies have estimated the capacity of consolidation to be one ite m at a time. Here we used a sequential-simultaneous manipulation to re-examine the limits of consolidating items into VSTM. Participants viewed briefly presen ted and masked color patches (targets), which were shown either sequentially or simultaneously. A probe color followed the targets and participants decided whether it matched one of the targets or was a novel color. In four experiments, we consistently found equal performance for sequential and simultaneous presentations for two targets. Worse performance in the simultaneous than the sequential condition was observed for larger set sizes (three and four). Contrary to previous results suggesting a severe capacity limit of one item, our results indicate that consolidation into VSTM can occur in parallel and without capacity limits for at least two items.
Selective attention to motion direction can modulate the strength of direction selective sensory responses regardless of their spatial locations. Although such spatially global modulation is thought to be a general property of feature-based attention, few studies have examined visual features other than motion. Here we used an adaptation protocol combined with attentional instructions to assess whether attention to orientation, a prominent feature in early visual processing, also exhibit such spatially global modulation. We adapted observers to an orientation by cueing them to attend to the orientation in a compound grating that was presented at a peripheral location. We then assessed the size of the tilt aftereffect at three locations that were never stimulated by the adapter. Attending to orientation produced a tilt aftereffect in these locations, indicating that attention modulated orientation selective mechanisms in remote locations from the adapter. Furthermore, there was no difference in the magnitude of the tilt aftereffect for test stimuli that were located at different distances and hemifields to the adapter. These results suggest that attention to orientation spreads uniformly across the visual field. Thus spatially global modulation seems to be a general property of feature-based attention and it provides a flexible mechanism to modulate feature salience across the visual field.
Sequential sampling models provide a useful framework for understanding human decision making. A key component of these models is an evidence accumulation process in which information is accrued over time to a threshold, at which point a choice is made. Previous neurophysiological studies on perceptual decision making have suggested accumulation occurs only in sensorimotor areas involved in making the action for the choice. Here we investigated the neural correlates of evidence accumulation in the human brain using functional magnetic resonance imaging (fMRI) while manipulating the quality of sensory evidence, the response modality, and the foreknowledge of the response modality. We trained subjects to perform a random dot motion direction discrimination task by either moving their eyes or pressing buttons to make their responses. In addition, they were cued about the response modality either in advance of the stimulus or after a delay. We isolated fMRI responses for perceptual decisions in both independently-defined sensorimotor areas and task-defined non-sensorimotor areas. We found neural signatures of evidence accumulation-a higher fMRI response on low coherence trials than high coherence trials-primarily in saccade-related sensorimotor areas (FEF, IPS) and non-sensorimotor areas in anterior insula and inferior frontal sulcus. Critically, such neural signatures did not depend on response modality or foreknowledge. These results help establish human brain areas involved in evidence accumulation and suggest that the neural mechanism for evidence accumulation is not specific to effectors. Instead, the neural system might accumulate evidence for particular stimulus features relevant to a perceptual task.
Human can flexibly attend to a variety of stimulus dimensions, including spatial location and various features such as color and direction of motion. While the locus of spatial attention has been hypothesized to be represented by priority maps encoded in several dorsal frontal and parietal areas, it is unknown how the brain represents attended features. Here we examined the distribution and organization of neural signals related to deployment of feature-based attention. Subjects viewed a compound stimulus containing two superimposed motion directions (or colors), and were instructed to perform an attention-demanding task on one of the directions (or colors). We found elevated and sustained fMRI response for the attention task compared to a neutral condition, without reliable differences in overall response amplitude between attending to different features. However, using multi-voxel pattern analysis, we were able to decode the attended feature in both early visual areas (V1 to hMT+) and frontal and parietal areas (e.g., IPS1-4 and FEF) that are commonly associated with spatial attention. Furthermore, analysis of the classifier weight maps showed that attending to motion and color evoked different patterns of activity, suggesting different neuronal subpopulations in these regions are recruited for attending to different feature dimensions. Thus, our finding suggests that rather than a purely spatial representation of priority, frontal and parietal cortical areas also contain multiplexed signals related to the priority of different non-spatial features.
Attending to a feature in one location can produce feature-specific modulation in a different location. This global feature-based attention effect has been demonstrated using two stimulus locations. Although the spread of feature-based attention is presumed to be constant across spatial locations, it has not been tested empirically. We examined the spread of feature-based attention by measuring attentional modulation of the motion aftereffect (MAE) at remote locations. Observers attended to one of two directions in a compound motion stimulus (adapter) and performed a speed-increment task. MAE was measured via a speed-nulling procedure for a test stimulus at different distances from the adapter. In Experiment 1, the adapter was at fixation, while the test stimulus was located at different eccentricities. We also measured the magnitude of baseline MAE for each location in two control conditions that did not require feature-based selection necessitated by a compound stimulus. In Experiment 2, the adapter and test stimuli were all located in the periphery at the same eccentricity. Our results showed that attention induced MAE spread completely across the visual field, indicating a genuine global effect. These results add to our understanding of the deployment of feature-based attention and provide empirical constraints on theories of visual attention.
Voluntary(endogenous, sustained) covert spatial attention selects relevant sensory information for prioritized processing. The behavioral and neural consequences of such selection have been extensively documented, but its phenomenologu has received little empirical investigation. We asked whether voluntary attention affects the subjective appearance of contrast- a fundamental dimension of visual perception. We used a demanding rapid serial visual presentation(RSVP) task to direct endogenous attention to a given location and measured perceived contrast at the attended and unattended locations. Attention increased perceived contrast of suprathreshold stimuli and also improved performance on a concurrent orientation discrimination task at the cued location. We ruled out response bias as an alternative account of the pattern of results. Thus, this study establishes that voluntary attention enhances perceived contrast. This phenomenological consequence links behavioral and neurophysiological studies on the effects of attention.
How does attention optimize our visual system for the task at hand? Two mechanisms have been proposed for how attention improves signal processing: gain and tuning. To distinguish between these two mechanisms we use the equivalent-noise paradigm, which measures performance as a function of external noise. In the present study we explored how spatial and feature-based attention affect performance by assessing their threshold-vs-noise (TvN) curves with regard to the signature behavioral effects of gain and tuning. Furthermore, we link our psychophysical results to neurophysiology by implementing a simple, biologically-plausible model to show that attention affects the gain and tuning of population responses differentially, depending on the type of attention being deployed: Whereas spatial attention operates by boosting the gain of the population response, feature-based attention operates by both boosting the gain and sharpening the tuning of the population response.
How humans recognize objects remains a contentious issue in current research on high-level vision. Here, I test the proposal by Wallis and Bulthoff (1999 Trends in Cognitive Sciences 3 22-31) suggesting that object representations can be learned through temporal association of multiple views of the same object. Participants first studied image sequences of novel, three-dimensional objects in a study block. On each trial, the images were from either an orderly sequence of depth-rotated views of the same object (SS), a scrambled sequence of those views (SR), or a sequence of different objects (RR). Recognition memory was assessed in a following test block. A within-object advantage was consistently observed--greater accuracy in the SR than the RR condition in all four experiments, greater accuracy in the SS than the RR condition in two experiments. Furthermore, spatiotemporal coherence did not produce better recognition than temporal coherence alone (similar or less accuracy in the SS compared to the SR condition). These results suggest that the visual system can use temporal regularity to build invariant object representations, via the temporal-association mechanism.
How does feature-based attention modulate neural responses? We used adaptation to quantify the effect of feature-based attention on orientation-selective responses in human visual cortex. Observers were adapted to two super-imposed oblique gratings while attending to one grating only. We measured the magnitude of attention-included orientation-selective adaptation both psychophysically, by the behavioral tilt aftereffect, and pschologically, using fMRI response adaptation. We found evidence for orientation-selective attentional modulation of neuronal responses- a lower fMRI response for the attended than the unattended orientation-in multiple visual areas, and a significant correlation between the magnitude of the tilt aftereffect and that of fMRI response adaptation in V1, the earliest site of orientation coding. These results show that feature-based attention can selectively increase the response of neuronal subpopulations that prefer the attended feature, even when the attended and unattended features are coded in the same visual areas and share the same retinotopic location.
We investigated the time course of feature-based attention and compared it to the time course of spatial attention in an experiment with identical stimuli and task. Observers detected a speed increment in a compound motion stimulus preceded by cues that indicated either the target location or direction. The cue-target stimulus-onset-asynchrony (SOA) was varied to assess the time course of the attentional effect. We found that spatial attention was deployed earlier than feature-based attention and that both types of attention improved performance to a similar extent at a longer SOA. Results indicate that attention is a flexible mechanism allowing us to efficiently select task-relevant information based on either spatial or feature dimensions, but that spatial attention exert its effects faster.
Human visual performance is better below than above fixation along the vertical meridian-a phenomenon we refer to as vertical meridian asymmetry (VMA). Here, we used fMRI to investigate the neural correlates of the VMA. We presented stimuli of two possible sizes and spatial frequencies on the horizontal and vertical meridians and analyzed the fMRI data in subregions of early visual cortex (V1/V2) that corresponded retinotopically to the stimulus locations. Asymmetries in both the spatial extent and amplitude of the fMRI measurements correlated with the behavioral VMA. These results demonstrate that the VMA has a neural basis at the earliest stages of cortical visual processing and imply that visual performance is limited by the pooled sensory responses of large populations of neurons in the visual cortex.
Selective attention enhances visual information processing, as measured by behavioral performance and neural activity. However, little is known about its effects on subjective experience. Here, we investigated the effect of transient (exogenous) attention on the appearance of visual motion, using a psychophysical procedure that directly measures appearance and controls for response bias. Observers viewed pairs of moving dot patterns and reported the motion direction of the more coherent pattern. Directing attention (via a peripheral precue) to a stimulus location increased its perceived coherence level and improved performance on a direction discrimination task. In a control experiment, we ruled out response bias by lengthening the time interval between the cue and the stimuli, so that the effect of transient attention could no longer be exerted. Our results are consistent with those of neurophysiological studies showing that attention modulates motion processing and provide evidence of a subjective perceptual correlate of attention, with a concomitant effect on performance.
The human visual system possesses a remarkable ability to reconstruct the shape of an object that is partly occluded by an interimposed surface. Behavioral results suggest that, under some circumstances, this perceptual process(termed amodal completion) progresses from an initial representation of local image features to a completed representation of a shape that may include features that are not explicitly present in the retinal image. Recent functional magnetic imaging(fMRI) studies have shown that the completed surface is represented in early visual cortical areas. We used fMRI adaptation, combined with brief, masked exposures, to track the amodal completion process as it unfolds in early visual cortical regions. We report evidence for an evolution of the neural representation from the image-based feature representation to the completed representation. Our method offers the possibility of measuring changes in cortical activity using fMRI over a time scale of a few hundred milliseconds.
When a visual stimulus suddenly appears, it captures attention, producing a transient improvement of performance on basic visual tasks. We investigate the effect of transient attention on stimulus representations in early visual areas using rapid event-related fMRI. Participants discriminated the orientation of one of two gratings preceded or followed by a non-predictive peripheral cue. Compared to control conditions, precueing the target location improved performance and produced a larger fMRI response in corresponding retinotopic areas. This enhancement progressively increased from striate to extrastriate areas. Control conditions indicated that the enhanced fMRI respose was not due to sensory summation of cue and target signals. Thus, an uninformative precue increases both perceptual performance and the concomitant stimulus-evoked activity in early visual areas. These results provide evidence regarding the retinotopically specific neural correlate for the effects of transient attention on early vision.
During apparent motion, spatially distinct items presented in alternation cause the perception of a visual stimulus smoothly traversing the intervening space where no physical stimulus exists. We used fMRI to determine whether the perceptual ‘filling-in’ that underlies this phenomenon has an early or late cortical locus. Subjects viewed a display comprised of concentric rings that elicited apparent motion (two concentric rings presented in alternation), flicker (the same rings presented simultaneously), or real motion. We independently localized the cortical regions corresponding to the path of apparent motion in early visual areas (V1, V2, VP, V3, V4v, V3A), as well as the human motion processing complex (MT+). Cortical activity in the path of apparent motion in early visual areas was similar in amplitude during both apparent motion and flicker. In contrast, cortical activity in MT+ was higher in amplitude during apparent motion than during flicker, but was lower in amplitude than during real motion. In addition, we observed overlap in the cortical loci of MT+ and the lateral occipital complex (LOC), a region involved in shape and object processing. This overlap suggests that these regions could directly interact and thereby support perceived object continuity during apparent motion.
Purpose: To describe retinotopic mapping of the visual cortex when a central scotoma is present. Design: Single observational case report. Methods: Scanning laser ophthalmoscope perimetry was used to define the site and stability of fixation and the area of dense scotoma. Functional magnetic resonance imaging of the visual cortex was performed while the patient viewed an expanding annular stimulus. Results: Retinotopic mapping of the visual cortex for a patient with a horseshoe scotoma from geographicatrophy involving the macular region showed a loss of stimulation to the cortical areas representing the site of the atrophic lesion. Conclusions: Cortical retinotopic mapping can be performed successfully in patients with central scotomas from macular disease. This study can serve as a basis for the future investigation of cortical plasticity in visual cortex. Ophthalmology 2004;111:1595–1598 © 2004 by the American Academy of Ophthalmology.
A network of fronto-parietal cortical areas is known to be involved in the control of visual attention, but the representational scope and specific function of these areas remains unclear. Recent neuro-imaging evidence has revealed the existence of both transient (attention-shift) and sustained (attention-maintenance) mechanisms of space-based and object-based attentional control. Here we investigate the neural mechanisms of feature-based attentional control in human cortex using rapid event-related functional magnetic resonance imaging (fMRI). Subjects viewed an aperture containing moving dots in which dot color and direction of motion changed once per second. At any given moment, observers attended to either motion or color. Two of six motion directions and two of six colors embedded in the stimulus stream cued subjects either to shift attention from the currently attended to the unattended feature or to maintain attention on the currently attended feature. Attentional modulation of the blood oxygenation level dependent (BOLD) fMRI signal was observed in early visual areas that are selective for motion and color. More importantly, both transient and sustained BOLD activity patterns were observed in different fronto-parietal cortical areas during shifts of attention. We suggest these differing temporal profiles reflect complementary roles in the control of attention to perceptual features.
Although both the object and the observer often move in natural environments, the effect of motion on visual object recognition has not been well documented. The authors examined the effect of a reversal in the direction of rotation on both explicit and implicit memory for novel, 3-dimensional objects. Participants viewed a series of continuously rotating objects and later made either an old–new recognition judgment or a symmetric–asymmetric decision. For both tasks, memory for rotating objects was impaired when the direction of rotation was reversed at test. These results demonstrate that dynamic information can play a role in visual object recognition and suggest that object representations can encode spatio-temporal information.
We argue that task requirements can be the determinant in generating different results in studies on visual object recognition. We investigated priming for novel visual objects in three implicit memory tasks. A study–test design was employed in which participants first viewed line drawings of unfamiliar objects and later made different decisions about structural aspects of the objects. Priming for both symmetric and asymmetric possible objects was observed in a task requiring a judgment of structural possibility. However, when the task was changed to one requiring a judgment of structural symmetry, only symmetric possible objects showed priming. Finally, in a matching task in which participants made a same–different judgment, only symmetric possible objects exhibited priming. These results suggest that an understanding of object representation will be most fruitful if it is based on careful analyses of both the task demands and their interaction(s) with encoding and retrieval processes.