Smooth pursuit deficits impact dynamic visual acuity in macular degeneration.

SIGNIFICANCE: Prior studies with large, highly visible targets report low smooth pursuit gains in individuals with macular degeneration (MD). We show that lower gains persist even when observers are pursuing a target that requires discrimination at the acuity limit. This low gain causes retinal slip, potentially leading to motion blur and target disappearance in the scotoma, which further compromise the visibility of moving object. PURPOSE: In this study, we examine whether the characteristics of smooth pursuit (pursuit gain and placement of the fixational locus relative to the target) change when the task requires dynamic visual acuity. METHODS: Using the scanning laser ophthalmoscope, we recorded smooth pursuit eye movements in 10 eyes of 6 MD participants and 7 eyes of 4 age-matched controls in response to leftward- or rightward-moving annular targets (O) that briefly (300 milliseconds) changed to a Landolt C at one of several time points during the pursuit trial. Participants were asked to pursue the target and indicate the direction of the C opening. RESULTS: Individuals with MD had lower pursuit gains and fewer saccades during the C presentation than during the O, compared with their age-matched peers. Further, pursuit gain, but not the distance of the retinal pursuit locus from the target, predicted task performance in the MD group. CONCLUSIONS: Our findings suggest that compromised pursuit gain in MD participants likely further compromises their dynamic visual acuity and thus ability to view moving targets.

Excitatory Contribution to Binocular Interactions in Human Visual Cortex Is Reduced in Strabismic Amblyopia.

Binocular summation in strabismic amblyopia is typically reported as being absent or greatly reduced in behavioral studies and is thought to be because of a preferential loss of excitatory interactions between the eyes. Here, we studied how excitatory and suppressive interactions contribute to binocular contrast interactions along the visual cortical hierarchy of humans with strabismic and anisometropic amblyopia in both sexes, using source-imaged steady-state visual evoked potentials (SSVEP) over a wide range of relative contrast between the two eyes. Dichoptic parallel grating stimuli modulated at unique temporal frequencies in each eye allowed us to quantify spectral response components associated with monocular inputs (self-terms) and the response components because of interaction of the inputs of the two eyes [intermodulation (IM) terms]. Although anisometropic amblyopes revealed a similar pattern of responses to normal-vision observers, strabismic amblyopes exhibited substantially reduced IM responses across cortical regions of interest (V1, V3a, hV4, hMT+ and lateral occipital cortex), indicating reduced interocular interactions in visual cortex. A contrast gain control model that simultaneously fits self- and IM-term responses within each cortical area revealed different patterns of binocular interactions between individuals with normal and disrupted binocularity. Our model fits show that in strabismic amblyopia, the excitatory contribution to binocular interactions is significantly reduced in both V1 and extra-striate cortex, whereas suppressive contributions remain intact. Our results provide robust electrophysiological evidence supporting the view that disruption of binocular interactions in strabismus or amblyopia is because of preferential loss of excitatory interactions between the eyes.SIGNIFICANCE STATEMENT We studied how excitatory and suppressive interactions contribute to binocular contrast interactions along the visual cortical hierarchy of humans with normal and amblyopic vision, using source-imaged SSVEP and frequency-domain analysis of dichoptic stimuli over a wide range of relative contrast between the two eyes. A dichoptic contrast gain control model was used to characterize these interactions in amblyopia and provided a quantitative comparison to normal vision. Our model fits revealed different patterns of binocular interactions between normal and amblyopic vision. Strabismic amblyopia significantly reduced excitatory contributions to binocular interactions, whereas suppressive contributions remained intact. Our results provide robust evidence supporting the view that the preferential loss of excitatory interactions disrupts binocular interactions in strabismic amblyopia.

Mapping residual stereopsis in macular degeneration.

Individuals with macular degeneration typically lose vision in the central region of one or both eyes. A binocular scotoma occurs when vision loss occurs in overlapping locations in both eyes, but stereopsis is impacted even in the non-overlapping region wherever the visual field in either eye is affected. We used a novel stereoperimetry protocol to measure local stereopsis across the visual field (up to 25° eccentricity) to determine how locations with functional stereopsis relate to the scotomata in the two eyes. Participants included those with monocular or binocular scotomata and age-matched controls with healthy vision. Targets (with or without depth information) were presented on a random dot background. Depth targets had true binocular disparity of 20' (crossed), whereas non-depth targets were defined by monocular cues such as contrast and dot density. Participants reported target location and whether it was in depth or flat. Local depth sensitivity (d') estimates were then combined to generate a stereopsis map. This stereopsis map was compared to the union of the monocular microperimetry estimates that mapped out the functional extent of the scotoma in each eye. The "union" prediction aligned with residual stereopsis, showing impaired stereopsis within this region and residual stereopsis outside this region. Importantly, the stereoblind region was typically more extensive than the binocular scotoma defined by the intersection (overlap) of the scotomata. This explains why individuals may have intact binocular visual fields but be severely compromised in tasks of daily living that benefit from stereopsis, such as eye-hand coordination and navigation.

Predicting Stereopsis in Macular Degeneration.

Each of our eyes sees a slightly different view of the physical world. Disparity is the small difference in position of features in the retinal images; stereopsis is the percept of depth from disparity. A distance between corresponding features in the retinal images of the two eyes smaller than the "upper disparity limit" yields a percept of depth; distances greater than this limit cause the two unfused monocular features to appear flattened into the fixation plane. This behavioral disparity limit is consistent with neurophysiological estimates of the largest disparity scale in primate, allowing us to relate physiological limits on plausible binocular interactions to separation between retinal locations. Here we test the hypothesis that this upper disparity limit predicts the presence of coarse stereopsis in humans with macular degeneration (MD), which affects the central retina but typically spares the periphery. The pattern of vision loss can be highly asymmetric, such that an intact location in one eye has a corresponding point in the other eye that lies within affected retina. Nevertheless, some individuals with MD have coarse stereopsis that is useful for eye-hand coordination. Our results show that individuals with MD (n = 25, male and female) have coarse stereopsis when the distance between intact retinal locations is less than the behavioral and physiological upper disparity limit at the corresponding eccentricity. Furthermore, for those without stereopsis, we can predict whether they can achieve stereopsis by using alternate retinal loci at further eccentricities whose separation is below the upper disparity limit.SIGNIFICANCE STATEMENT We show that the largest separation between features in the two eyes that yields a percept of depth in humans is related to the largest disparity scale in macaque medial temporal area and to the estimated size of the receptive fields in human depth-sensitive cortical regions. This upper disparity limit also predicts whether individuals with retinal damage due to macular degeneration will have stereopsis. Individuals have stereopsis when the separation between intact retinal locations in the two eyes is smaller than the upper disparity limit measured behaviorally. Our results indicate the importance of the behavioral upper disparity limit as a predictor for stereopsis in populations with retinal damage.

Contrast Normalization Accounts for Binocular Interactions in Human Striate and Extra-striate Visual Cortex.

During binocular viewing, visual inputs from the two eyes interact at the level of visual cortex. Here we studied binocular interactions in human visual cortex, including both sexes, using source-imaged steady-state visual evoked potentials over a wide range of relative contrast between two eyes. The ROIs included areas V1, V3a, hV4, hMT+, and lateral occipital cortex. Dichoptic parallel grating stimuli in each eye modulated at distinct temporal frequencies allowed us to quantify spectral components associated with the individual stimuli from monocular inputs (self-terms) and responses due to interaction between the inputs from the two eyes (intermodulation [IM] terms). Data with self-terms revealed an interocular suppression effect, in which the responses to the stimulus in one eye were reduced when a stimulus was presented simultaneously to the other eye. The suppression magnitude varied depending on visual area, and the relative contrast between the two eyes. Suppression was strongest in V1 and V3a (50% reduction) and was least in lateral occipital cortex (20% reduction). Data with IM terms revealed another form of binocular interaction, compared with self-terms. IM response was strongest at V1 and was least in hV4. Fits of a family of divisive gain control models to both self- and IM-term responses within each cortical area indicated that both forms of binocular interaction shared a common gain control nonlinearity. However, our model fits revealed different patterns of binocular interaction along the cortical hierarchy, particularly in terms of excitatory and suppressive contributions.SIGNIFICANCE STATEMENT Using source-imaged steady-state visual evoked potentials and frequency-domain analysis of dichoptic stimuli, we measured two forms of binocular interactions: one is associated with the individual stimuli that represent interocular suppression from each eye, and the other is a direct measure of interocular interaction between inputs from the two eyes. We demonstrated that both forms of binocular interactions share a common gain control mechanism in striate and extra-striate cortex. Furthermore, our model fits revealed different patterns of binocular interaction along the visual cortical hierarchy, particularly in terms of excitatory and suppressive contributions.

Mapping the binocular scotoma in macular degeneration.

When the scotoma is binocular in macular degeneration (MD), it often obscures objects of interest, causing individuals to miss information. To map the binocular scotoma as precisely as current methods that map the monocular scotoma, we propose an iterative eye-tracker method. Study participants included nine individuals with MD and four age-matched controls. We measured the extent of the monocular scotomata using a scanning laser ophthalmoscope/optical coherence tomography (SLO/OCT). Then, we precisely mapped monocular and binocular scotomata with an eye tracker, while fixation was monitored. Participants responded whenever they detected briefly flashed dots, which were first presented on a coarse grid, and then at manually selected points to refine the shape and edges of the scotoma. Monocular scotomata measured in the SLO and eye tracker are highly similar, validating the eye-tracking method for scotoma mapping. Moreover, all participants used clustered fixation loci corresponding to their dominant preferred fixation locus. Critically, for individuals with binocular scotomata, the binocular map from the eye tracker was consistent with the overlap of the monocular scotoma profiles from the SLO. Thus, eye-tracker-based perimetry offers a reliable and sensitive tool for measuring both monocular and binocular scotomata, unlike the SLO/OCT that is limited to monocular viewing.

Motion perception in central field loss.

Motion information is essential in daily life because it provides cues to depth, timing, object identification, and self-motion, as well as input to the oculomotor system. As the peripheral visual field is exquisitely sensitive to motion, we investigated the periphery of individuals with central visual field loss (CFL) to determine whether speed and direction discrimination are intact in this population. We compared CFL participants' (N = 8), older (N = 6), and young controls' (N = 6) ability to discriminate motion speed and direction in a two-spatial-alternative forced-choice design. Participants viewed moving dots on the left and right of a fixation marker and judged which side had the faster speed or more clockwise direction. For the young control group, we repeated the experiment with the stimulus limited to thin strips of fixed width at eccentricities of 5°, 10°, and 15°. There was no significant difference in mean speed or direction discrimination thresholds of CFL participants and older controls for either velocity. Young controls had significantly lower thresholds than the CFL group for both tasks. We did not find an effect of visual acuity, viewing eccentricity, or scotoma location on individuals' ability to discriminate speed or direction. Our results indicate that for high-visibility stimuli moving at 5°-10°/s, speed and direction discrimination are intact in the periphery of individuals with CFL.

The upper disparity limit increases gradually with eccentricity.

Stereopsis is important for tasks of daily living such as eye-hand coordination. It is best in central vision but is also mediated by the periphery. Previously we have shown that individuals with central-field loss who have residual stereopsis in the periphery perform better at an eye-hand-coordination task when they perform the task binocularly rather than monocularly. Here we seek to determine what sets the limit of stereopsis, defined as the largest disparity that supports the sustained appearance of depth, in the near periphery in healthy individuals. While stereoacuity thresholds increase sharply with eccentricity, Panum's area increases much more slowly. We used a rigorous method to determine the uppermost limit of disparity. At long durations, the two half-images that define a large disparity appear as two isolated targets in the same flat plane; small incremental changes in disparity produce changes in the separation between the half-images, and disparity magnitude can be judged on the basis of separation, like a monocular width judgment. The disparity limit is the point at which the threshold for judging dichoptic separation between the half-images is equal to the monocular width-discrimination threshold. The disparity limit at 10° was a factor of 2-4 times larger than the fovea, regardless of the meridian tested. The increase in the disparity limit with eccentricity was shallow, similar to that of Panum's area. Within this disparity limit, disparity increment thresholds were comparable for foveal and peripheral targets, illustrating the significance and utility of peripheral stereopsis, especially in the absence of foveal stereopsis.

The spatial and temporal properties of attentional selectivity for saccades and reaches.

The preparation and execution of saccades and goal-directed movements elicits an accompanying shift in attention at the locus of the impending movement. However, some key aspects of the spatiotemporal profile of this attentional shift between eye and hand movements are not resolved. While there is evidence that attention is improved at the target location when making a reach, it is not clear how attention shifts over space and time around the movement target as a saccade and a reach are made to that target. Determining this spread of attention is an important aspect in understanding how attentional resources are used in relation to movement planning and guidance in real world tasks. We compared performance on a perceptual discrimination paradigm during a saccade-alone task, reach-alone task, and a saccade-plus-reach task to map the temporal profile of the premotor attentional shift at the goal of the movement and at three surrounding locations. We measured performance relative to a valid baseline level to determine whether motor planning induces additional attentional facilitation compared to mere covert attention. Sensitivity increased relative to movement onset at the target and at the surrounding locations, for both the saccade-alone and saccade-plus-reach conditions. The results suggest that the temporal profile of the attentional shift is similar for the two tasks involving saccades (saccade-alone and saccade-plus-reach tasks), but is very different when the influence of the saccade is removed. In this case, performance in the saccade-plus-reach task reflects the lower sensitivity observed when a reach-alone task is being conducted. In addition, the spatial profile of this spread of attention is not symmetrical around the target. This suggests that when a saccade and reach are being planned together, the saccade drives the attentional shift, and the reach-alone carries little attentional weight.

Attention to Multiple Objects Facilitates Their Integration in Prefrontal and Parietal Cortex.

Selective attention is known to interact with perceptual organization. In visual scenes, individual objects that are distinct and discriminable may occur on their own, or in groups such as a stack of books. The main objective of this study is to probe the neural interaction that occurs between individual objects when attention is directed toward one or more objects. Here we record steady-state visual evoked potentials via electrocorticography to directly assess the responses to individual stimuli and to their interaction. When human participants attend to two adjacent stimuli, prefrontal and parietal cortex shows a selective enhancement of only the neural interaction between stimuli, but not the responses to individual stimuli. When only one stimulus is attended, the neural response to that stimulus is selectively enhanced in prefrontal and parietal cortex. In contrast, early visual areas generally manifest responses to individual stimuli and to their interaction regardless of attentional task, although a subset of the responses is modulated similarly to prefrontal and parietal cortex. Thus, the neural representation of the visual scene as one progresses up the cortical hierarchy becomes more highly task-specific and represents either individual stimuli or their interaction, depending on the behavioral goal. Attention to multiple objects facilitates an integration of objects akin to perceptual grouping.SIGNIFICANCE STATEMENT Individual objects in a visual scene are seen as distinct entities or as parts of a whole. Here we examine how attention to multiple objects affects their neural representation. Previous studies measured single-cell or fMRI responses and obtained only aggregate measures that combined the activity to individual stimuli as well as their potential interaction. Here, we directly measure electrocorticographic steady-state responses corresponding to individual objects and to their interaction using a frequency-tagging technique. Attention to two stimuli increases the interaction component that is a hallmark for perceptual integration of stimuli. Furthermore, this stimulus-specific interaction is represented in prefrontal and parietal cortex in a task-dependent manner.

Texture segmentation influences the spatial profile of presaccadic attention.

Attention is important for selecting targets for action. Several studies have shown that attentional selection precedes eye movements to a target, and results in an enhanced sensitivity at the saccade goal. Typically these studies have used isolated targets on blank backgrounds, which are rare in real-world situations. Here, we examine the spatial profile of sensitivity around a saccade target on a textured background and how the influence of the surrounding context develops over time. We used two textured backgrounds: a uniform texture, and a concentric arrangement of an inner and an outer texture with orthogonal orientations. For comparison, we also measured sensitivity around the target on a blank background. The spatial profile of sensitivity was measured with a brief, dim, probe flashed around the saccade target. When the target was on a blank or a uniformly textured background, spatial sensitivity peaked near the target location around 350 ms after cue onset and declined with distance from the target. However, when the background was made up of an inner and outer texture, sensitivity to the inner texture was uniformly high, peaking at about 350 ms after cue onset, suggesting that the entire inner texture was selected along with the target. The enhancement of sensitivity on the inner texture was much smaller when observers attended the target covertly and performed the probe-detection task. Thus, our results suggest that the surface representation around the target is taken into account when an observer actively plans to interact with the target.

Cortical sources of Vernier acuity in the human visual system: An EEG-source imaging study.

Vernier acuity determines the relative position of visual features with a precision better than the sampling resolution of cone receptors in the retina. Because Vernier displacement is thought to be mediated by orientation-tuned mechanisms, Vernier acuity is presumed to be processed in striate visual cortex (V1). However, there is considerable evidence suggesting that Vernier acuity is dependent not only on structures in V1 but also on processing in extrastriate cortical regions. Here we used functional magnetic resonance imaging-informed electroencephalogram source imaging to localize the cortical sources of Vernier acuity in observers with normal vision. We measured suprathreshold and near-threshold responses to Vernier onset/offset stimuli at different stages of the visual cortical hierarchy, including V1, hV4, lateral occipital cortex (LOC), and middle temporal cortex (hMT+). These responses were compared with responses to grating on/off stimuli, as well as to stimuli that control for lateral motion in the Vernier task. Our results show that all visual cortical regions of interest (ROIs) responded to both suprathreshold Vernier and grating stimuli. However, thresholds for Vernier displacement (Vernier acuity) were lowest in V1 and LOC compared with hV4 and hMT+, whereas all visual ROIs had identical thresholds for spatial frequency (grating acuity) and for relative motion. The cortical selectivity of sensitivity to Vernier displacement provides strong evidence that LOC, in addition to V1, is involved in Vernier acuity processing. The robust activation of LOC might be related to the sensitivity to the relative position of features, which is common to Vernier displacement and to some kinds of texture segmentation.

Training eye movements for visual search in individuals with macular degeneration.

We report a method to train individuals with central field loss due to macular degeneration to improve the efficiency of visual search. Our method requires participants to make a same/different judgment on two simple silhouettes. One silhouette is presented in an area that falls within the binocular scotoma while they are fixating the center of the screen with their preferred retinal locus (PRL); the other silhouette is presented diametrically opposite within the intact visual field. Over the course of 480 trials (approximately 6 hr), we gradually reduced the amount of time that participants have to make a saccade and judge the similarity of stimuli. This requires that they direct their PRL first toward the stimulus that is initially hidden behind the scotoma. Results from nine participants show that all participants could complete the task faster with training without sacrificing accuracy on the same/different judgment task. Although a majority of participants were able to direct their PRL toward the initially hidden stimulus, the ability to do so varied between participants. Specifically, six of nine participants made faster saccades with training. A smaller set (four of nine) made accurate saccades inside or close to the target area and retained this strategy 2 to 3 months after training. Subjective reports suggest that training increased awareness of the scotoma location for some individuals. However, training did not transfer to a different visual search task. Nevertheless, our study suggests that increasing scotoma awareness and training participants to look toward their scotoma may help them acquire missing information.

Accuracy of eye position for saccades and smooth pursuit.

In this study, we address the question of whether a target is foveated during smooth pursuit. Specifically, we examine whether smooth pursuit eye movements land near the center-of-mass of the target, as is the case for saccades. To that end, we instructed eight untrained, healthy participants to follow moving targets, presented monocularly in a scanning laser ophthalmoscope. Stimuli moved either in a modified step-ramp (smooth pursuit), or made a single step (saccade), stepping 6° from the center. Targets were ring-shaped and either 0.6° or 1.7° in diameter. In an additional set of experiments, two participants collected more extensive data on smooth pursuit and saccades for a larger range of target sizes (0.6°, 1.7°, or 4.3°). During pursuit, eyes were rarely placed at target center, even when participants' fixational stability was taken into account. Furthermore, there was a clear tendency for distance from target center to increase with target size. This outcome was in contrast to saccades, where there was no effect of target size across participants. The difference in foveal placement between the two types of eye movements is consistent with their different purposes: closer inspection of the target for saccades versus maintenance of the target in the visual field for smooth pursuit.

Smooth pursuit eye movements in patients with macular degeneration.

Currently, there are no quantitative studies of smooth pursuit, a behavior attributed to the fovea, in individuals with macular degeneration (MD). We hypothesize that pursuit in MD patients depends on the relative positions of the scotoma and target trajectory. We tested this hypothesis with a scanning laser ophthalmoscope (SLO), which allows for direct visualization of the target on the damaged retina. Monocular microperimetry and eye movements were assessed in eleven individuals with differing degrees of MD. Observers were asked to visually track a 1.7° target that moved in one of eight radial directions at 5°/s-6°/s. Consistent with our hypothesis, pursuit metrics depended on whether the target moved into or out of scotoma. Pursuit gains decreased with increasing scotoma extent in the target's heading direction (p = 0.017). Latencies were higher when the scotoma was present along the target trajectory (in either starting or heading directions, p < 0.001). Furthermore, an analysis of retinal position shows that targets fell on the fixational locus nearly 50% of the time. The results suggest that MD patients are capable of smooth pursuit eye movements, but are limited by target trajectory and scotoma characteristics.

Degraded attentional modulation of cortical neural populations in strabismic amblyopia.

Behavioral studies have reported reduced spatial attention in amblyopia, a developmental disorder of spatial vision. However, the neural populations in the visual cortex linked with these behavioral spatial attention deficits have not been identified. Here, we use functional MRI-informed electroencephalography source imaging to measure the effect of attention on neural population activity in the visual cortex of human adult strabismic amblyopes who were stereoblind. We show that compared with controls, the modulatory effects of selective visual attention on the input from the amblyopic eye are substantially reduced in the primary visual cortex (V1) as well as in extrastriate visual areas hV4 and hMT+. Degraded attentional modulation is also found in the normal-acuity fellow eye in areas hV4 and hMT+ but not in V1. These results provide electrophysiological evidence that abnormal binocular input during a developmental critical period may impact cortical connections between the visual cortex and higher level cortices beyond the known amblyopic losses in V1 and V2, suggesting that a deficit of attentional modulation in the visual cortex is an important component of the functional impairment in amblyopia. Furthermore, we find that degraded attentional modulation in V1 is correlated with the magnitude of interocular suppression and the depth of amblyopia. These results support the view that the visual suppression often seen in strabismic amblyopia might be a form of attentional neglect of the visual input to the amblyopic eye.

Stop before you saccade: Looking into an artificial peripheral scotoma.

We investigated whether adults with healthy vision can move their eyes toward an informative target area that is initially hidden by a gaze-contingent scotoma in the periphery when they are under time pressure. In the experimental task, participants had to perform an object-comparison task requiring a same-different judgment about two silhouettes. One silhouette was visible, whereas the other was hidden under the scotoma. Despite time pressure and the presence of the visible silhouette, most participants were able to move their eyes toward the informative region to reveal the hidden silhouette. Saccades to the hidden stimulus occurred when the visible stimulus was presented directly opposite in either fixed or variable locations and when the visible stimulus was presented at an adjacent location. Older participants were also able to perform this task. First saccades in the direction of the hidden stimulus had longer latencies compared with saccades toward the visible stimulus. This suggests the use of a deliberate, nonreflexive saccade strategy ("stop before you saccade"). A subset of participants occasionally made curved saccades that were aimed first toward the visible stimulus and then toward the hidden stimulus. We discuss the implications of our findings for patients who have a biological scotoma, for example, in macular degeneration.

The influence of segmentation and uncertainty on target selection.

This study examines how two factors affect target selection: the contiguity of the target with the surrounding surface and certainty about target location. Previous studies indicate that a target among distractors is easier to find when the search items are on the same surface rather than different surfaces. In contrast, our recent study indicates that when the target is in a known location, sensitivity to the target is higher when it is clearly separated from the surrounding surface. Here we examine the effects of both surface contiguity and uncertainty about target location on contrast discrimination. Observers were asked to detect a contrast change on a grating target that was either segmented or contiguous with the surround grating and occurred either at a known or unknown location. Thresholds for contrast discrimination depended critically on both segmentation and location uncertainty. When the contrast change appeared at a known location isolated from the background, segmentation aided the selection of the target location, but when the contrast change occurred at an unknown location on a contiguous background, grouping of the surface as a single entity aided the detection of the target location as a discontinuity from the surface.

The selectivity of task-dependent attention varies with surrounding context.

Attention is thought to operate by enhancing the target of interest and suppressing the surroundings. We hypothesized that the spatial profile of attention depends on the surround's relationship to the target. Using high-density electroencephalographic measurements, we examined the spatial profile of attention to a grating target surrounded by an annular grating that was either coextensive with the target (unsegmented) or appeared segmented from it due to a gap or phase offset. We directly probed the spread of attention from the central target into the surround by flickering the surround and monitoring frequency-tagged steady-state visual-evoked potentials. Observers were required to detect a contrast increment that occurred only on the target. Successful detection of the increment required selecting the target and suppressing the surround, particularly when the target did not readily segment from the surround. The profile of attention was investigated in five visual regions of interest (ROIs) (V1, V4, V3A, lateral occipital complex, and human middle temporal area), mapped in a separate anatomical magnetic resonance imaging scan. We found that in most ROIs, attention to the target generated smaller responses from the surrounding annulus when it was contiguous compared with when it was clearly segmented. This result shows that the profile of attention depends on task demands and on surrounding context; attention is tightly focused when the target region needs to be isolated but loosely focused when the target region is clearly segmented.

Attention selects informative neural populations in human V1.

In a neural population driven by a simple grating stimulus, different subpopulations are maximally informative about changes to the grating's orientation and contrast. In theory, observers should attend to the optimal subpopulation when switching between orientation and contrast discrimination tasks. Here we used source-imaged, steady-state visual evoked potentials and visual psychophysics to determine whether this is the case. Observers fixated centrally while static targets were presented bilaterally along with a cue indicating task type (contrast or orientation modulation detection) and task location (left or right). Changes in neuronal activity were measured by quantifying frequency-tagged responses from flickering "reporter" gratings surrounding the targets. To determine the orientation tuning of attentionally modulated neurons, we measured responses for three different probe-reporter angles: 0, 20, and 45°. We estimated frequency-tagged cortical activity using a minimum norm inverse procedure combined with realistic MR-derived head models and retinotopically mapped visual areas. Estimates of neural activity from regions of interest centered on V1 showed that attention to a spatial location clearly increased the amplitude of the neural response in that location. More importantly, the pattern of modulation depended on the task. For orientation discrimination, attentional modulation showed a sharp peak in the population tuned 20° from the target orientation, whereas for contrast discrimination the enhancement was more broadly tuned. Similar tuning functions for orientation and contrast discrimination were obtained from psychophysical adaptation studies. These findings indicate that humans attend selectively to the most informative neural population and that these populations change depending on the nature of the task.