Functional MRI of visual cortex predicts training-induced recovery in stroke patients with homonymous visual field defects.

Post-chiasmatic damage to the visual system leads to homonymous visual field defects (HVDs), which can severely interfere with daily life activities. Visual Restitution Training (VRT) can recover parts of the affected visual field in patients with chronic HVDs, but training outcome is variable. An untested hypothesis suggests that training potential may be largest in regions with 'neural reserve', where cortical responses to visual stimulation do not lead to visual awareness as assessed by Humphrey perimetry-a standard behavioural visual field test. Here, we tested this hypothesis in a sample of twenty-seven hemianopic stroke patients, who participated in an assiduous 80-hour VRT program. For each patient, we collected Humphrey perimetry and wide-field fMRI-based retinotopic mapping data prior to training. In addition, we used Goal Attainment Scaling to assess whether personal activities in daily living improved. After training, we assessed with a second Humphrey perimetry measurement whether the visual field was improved and evaluated which personal goals were attained. Confirming the hypothesis, we found significantly larger improvements of visual sensitivity at field locations with neural reserve. These visual field improvements implicated both regions in primary visual cortex and higher order visual areas. In addition, improvement in daily life activities correlated with the extent of visual field enlargement. Our findings are an important step toward understanding the mechanisms of visual restitution as well as predicting training efficacy in stroke patients with chronic hemianopia.

Impaired visual competition in patients with homonymous visual field defects.

Intense visual training can lead to partial recovery of visual field defects caused by lesions of the primary visual cortex. However, the standard visual detection and discrimination tasks, used to assess this recovery process tend to ignore the complexity of the natural visual environment, where multiple stimuli continuously interact. Visual competition is an essential component for natural search tasks and detecting unexpected events. Our study focused on visual decision-making and to what extent the recovered visual field can compete for attention with the 'intact' visual field. Nine patients with visual field defects who had previously received visual discrimination training, were compared to healthy age-matched controls using a saccade target-selection paradigm, in which participants actively make a saccade towards the brighter of two flashed targets. To further investigate the nature of competition (feed-forward or feedback inhibition), we presented two flashes that reversed their intensity difference during the flash. Both competition between recovered visual field and intact visual field, as well as competition within the intact visual field, were assessed. Healthy controls showed the expected primacy effect; they preferred the initially brighter target. Surprisingly, choice behaviour, even in the patients' supposedly 'intact' visual field, was significantly different from the control group for all but one. In the latter patient, competition was comparable to the controls. All other patients showed a significantly reduced preference to the brighter target, but still showed a small hint of primacy in the reversal conditions. The present results indicate that patients and controls have similar decision-making mechanisms but patients' choices are affected by a strong tendency to guess, even in the intact visual field. This tendency likely reveals slower integration of information, paired with a lower threshold. Current rehabilitation should therefore also include training focused on improving visual decision-making of the defective and the intact visual field.

Visual perception of axes of head rotation.

Registration of ego-motion is important to accurately navigate through space. Movements of the head and eye relative to space are registered through the vestibular system and optical flow, respectively. Here, we address three questions concerning the visual registration of self-rotation. (1) Eye-in-head movements provide a link between the motion signals received by sensors in the moving eye and sensors in the moving head. How are these signals combined into an ego-rotation percept? We combined optic flow of simulated forward and rotational motion of the eye with different levels of eye-in-head rotation for a stationary head. We dissociated simulated gaze rotation and head rotation by different levels of eye-in-head pursuit. We found that perceived rotation matches simulated head- not gaze-rotation. This rejects a model for perceived self-rotation that relies on the rotation of the gaze line. Rather, eye-in-head signals serve to transform the optic flow's rotation information, that specifies rotation of the scene relative to the eye, into a rotation relative to the head. This suggests that transformed visual self-rotation signals may combine with vestibular signals. (2) Do transformed visual self-rotation signals reflect the arrangement of the semi-circular canals (SCC)? Previously, we found sub-regions within MST and V6(+) that respond to the speed of the simulated head rotation. Here, we re-analyzed those Blood oxygenated level-dependent (BOLD) signals for the presence of a spatial dissociation related to the axes of visually simulated head rotation, such as have been found in sub-cortical regions of various animals. Contrary, we found a rather uniform BOLD response to simulated rotation along the three SCC axes. (3) We investigated if subject's sensitivity to the direction of the head rotation axis shows SCC axes specifcity. We found that sensitivity to head rotation is rather uniformly distributed, suggesting that in human cortex, visuo-vestibular integration is not arranged into the SCC frame.

Transfer effects of training-induced visual field recovery in patients with chronic stroke.

OBJECTIVE: Visual training of light detection in the transition zone between blind and healthy hemianopic visual fields leads to improvement of color and simple pattern recognition. Recently, we demonstrated that visual field enlargement (VFE) also occurs when an area just beyond the transition zone is stimulated. In the current study, we attempted to determine whether this peripheral training also causes improvement in color and shape perception and reading speed. Further, we evaluated which measure of VFE relates best to improvements in performance: the average border shift (ABS) in degrees or the estimated amount of cortical surface gain (ECSG) in millimeters, using the cortical magnification factor (CMF). METHOD: Twelve patients received 40 sessions of 1-hour restorative function training (RFT). Before and after training, we measured visual fields and reading speed. Additionally, color and shape perception in the trained visual field area was measured in 7 patients. RESULTS: VFE was found for 9 of 12 patients. Significant improvements were observed in reading speed for 8 of 12 patients and in color and shape perception for 3 of 7 patients. ECSG correlates significantly with performance; ABS does not. Our data indicate that the threshold ECSG, needed for significant changes in color and shape perception and reading speed, is about 6 mm. CONCLUSIONS: White stimulus training-induced VFE can lead to improved color and shape perception and to increased reading speed in and beyond the pretraining transition zone if ECSG is sufficiently large. The latter depends on the eccentricity of the VFE.

Oculomotor behavior of hemianopic chronic stroke patients in a driving simulator is modulated by vision training.

BACKGROUND: Visual Restorative function training aims to decrease visual field defect size after acquired brain damage. Some chronic stroke patients regain permission to drive a car after training. This points to a concomitant change in oculomotor behavior, because visual field enlargement is hardly ever large enough for legal driving. This study investigated vRFT-induced changes in oculomotor behavior, using a driving simulator. METHODS: Driving performance and oculomotor behavior were measured before and after training in 6 hemianopia patients who had trained 65 hours with vRFT on a PC at home. RESULTS: Two patients showed negligible visual field enlargement (VFE) and four showed moderate to substantial VFE. Because less visual cortex is devoted to the processing of peripheral than central visual field the same VFE corresponds to less functional restoration of cortex when the defect is at high eccentricity. When this is taken into account, then precisely the two patients that showed the largest cortical gains made significantly more eye movements in the direction of their visual field defect after training. CONCLUSIONS: vRFT with mandatory eye fixation can result in increased eye movement behavior towards the defect. Our study suggests that a threshold amount of cortical functional restoration is required for this effect.

Effects of vision restoration training on early visual cortex in patients with cerebral blindness investigated with functional magnetic resonance imaging.

Cerebral blindness is a loss of vision as a result of postchiasmatic damage to the visual pathways. Parts of the lost visual field can be restored through training. However, the neuronal mechanisms through which training effects occur are still unclear. We therefore assessed training-induced changes in brain function in eight patients with cerebral blindness. Visual fields were measured with perimetry and retinotopic maps were acquired with functional magnetic resonance imaging (fMRI) before and after vision restoration training. We assessed differences in hemodynamic responses between sessions that represented changes in amplitudes of neural responses and changes in receptive field locations and sizes. Perimetry results showed highly varied visual field recovery with shifts of the central visual field border ranging between 1 and 7°. fMRI results showed that, although retinotopic maps were mostly stable over sessions, there was a small shift of receptive field locations toward a higher eccentricity after training in addition to increases in receptive field sizes. In patients with bilateral brain activation, these effects were stronger in the affected than in the intact hemisphere. Changes in receptive field size and location could account for limited visual field recovery (± 1°), although it could not account for the large increases in visual field size that were observed in some patients. Furthermore, the retinotopic maps strongly matched perimetry measurements before training. These results are taken to indicate that local visual field enlargements are caused by receptive field changes in early visual cortex, whereas large-scale improvement cannot be explained by this mechanism.

Intermittent ambiguous stimuli: implicit memory causes periodic perceptual alternations.

When viewing a stimulus that has multiple plausible real-world interpretations, perception alternates between these interpretations every few seconds. Alternations can be halted by intermittently removing the stimulus from view. The same interpretation dominates over many successive presentations, and perception stabilizes. Here we study perception during long sessions of such intermittent presentation. We demonstrate that, rather than causing truly stable perception, intermittent presentation gives rise to a perceptual alternation cycle with its own characteristics and dependencies, different from those during continuous presentation. Alternations during intermittent viewing typically occur once every few minutes--much less frequently than the seconds-scale alternations during continuous viewing. Strikingly, alternations during intermittent viewing occur at fairly regular intervals, making for a surprisingly periodic alternation cycle. The duration of this cycle becomes longer as the blank duration between presentations is increased, reaching dozens of minutes in some cases. We interpret our findings in terms of a mathematical model that describes a neural network with competition between alternative interpretations. Network sensitivities depend on prior dominance, thus providing a memory for past perception. Slow changes in sensitivity produce both perceptual stabilization and the regular but infrequent alternations, meaning that the same memory traces are responsible for both. This model provides a good description of psychophysical findings, and offers several indications regarding their neural basis.

Direct extraction of curvature-based metric shape from stereo by view-modulated receptive fields.

Any computation of metric surface structure from horizontal disparities depends on the viewing geometry, and analysing this dependence allows us to narrow down the choice of viable schemes. For example, all depth-based or slant-based schemes (i.e. nearly all existing models) are found to be unrealistically sensitive to natural errors in vergence. Curvature-based schemes avoid these problems and require only moderate, more robust view-dependent corrections to yield local object shape, without any depth coding. This fits the fact that humans are strikingly insensitive to global depth but accurate in discriminating surface curvature. The latter also excludes coding only affine structure. In view of new adaptation results, our goal becomes to directly extract retinotopic fields of metric surface curvatures (i.e. avoiding intermediate disparity curvature). To find a robust neural realisation, we combine new exact analysis with basic neural and psychophysical constraints. Systematic, step-by-step 'design' leads to neural operators which employ a novel family of 'dynamic' receptive fields (RFs), tuned to specific (bi-)local disparity structure. The required RF family is dictated by the non-Euclidean geometry that we identify as inherent in cyclopean vision. The dynamic RF-subfield patterns are controlled via gain modulation by binocular vergence and version, and parameterised by a cell-specific tuning to slant. Our full characterisation of the neural operators invites a range of new neurophysiological tests. Regarding shape perception, the model inverts widely accepted interpretations: It predicts the various types of errors that have often been mistaken for evidence against metric shape extraction.

Attentional control over either of the two competing percepts of ambiguous stimuli revealed by a two-parameter analysis: means do not make the difference.

We studied distributions of perceptual rivalry reversals, as defined by the two fitted parameters of the Gamma distribution. We did so for a variety of bi-stable stimuli and voluntary control exertion tasks. Subjects' distributions differed from one another for a particular stimulus and control task in a systematic way that reflects a constraint on the describing parameters. We found a variety of two-parameter effects, the most important one being that distributions of subjects differ from one another in the same systematic way across different stimuli and control tasks (i.e., a fast switcher remains fast across all conditions in a parameter-specified way). The cardinal component of subject-dependent variation was not the conventionally used mean reversal rate, but a component that was oriented-for all stimuli and tasks-roughly perpendicular to the mean rate. For the Necker cube, we performed additional experiments employing specific variations in control exertion, suggesting that subjects have to a considerable extent independent control over the reversal rate of either of the two competing percepts.

Collision judgment of objects approaching the head.

Recent investigations have indicated that human perception of the trajectory of objects approaching in the horizontal plane is precise but biased away from straight ahead. This is remarkable because it could mean that subjects perceive objects that approach on a collision course as missing the head. Approach within the horizontal plane through the eyes and the fixation point (the plane of regard) is special, as general motions will also have a component of motion perpendicular to the plane of regard. Thus, we investigated three-dimensional motion perception in the vicinity of the head, including vertical components. Subjects judged whether an object that moved in the mid-sagittal plane was going to hit below or above a well-known reference point on the face like the center of the chin or the forehead (perceptual task). Tactile and proprioceptive information about the reference point significantly improved precision. Precision did not change with distance of the approaching target or with fixation direction. Bias was virtually absent for these vertical motions. When subjects pointed with their index finger to the perceived location of impact on their face (visuo-motor task), they overestimated (1.7 cm) the horizontal eccentricity of the point of impact (pointing task). Vertical bias, however, was again virtually absent. Interestingly, when trajectories intersected the plane of regard, higher precision was observed in the perceptual task regardless of the other conditions. In contrast, neither bias nor precision of the pointing task changed significantly when the trajectories intersected the plane of regard. When asked to point to the location where a trajectory intersected the plane of regard, subjects overestimated the depth component of this intersection location by about 3 cm. The absence of perceptual and pointing bias in the vertical direction in contrast to the clear horizontal bias suggests that different (combinations of) cues are used to judge these components of the trajectory of an approaching object. The results of our perceptual task suggest a role for somatosensory signals in the visual judgment of impending impact.

An invariant for timing of saccades during visual search.

The variable latency of a saccade to the onset of a single target reveals our brain's hypothesis testing about the target's presence. Search in complex scenes involves multiple objects that compete to become fixated. The initiation of a saccade in this case involves two hypotheses: (1) a potential target is present outside the fovea and (2) the currently fixated object is not the target. Previous models suggest that these hypotheses are evaluated independently, each involving a decision signal that races towards threshold. We show here that the skewed latency distributions during search comply with strong competition between these decision signals rather than independence. Moreover, the thresholds for the two competing processes are not independent either but conform to an invariant that suggests that saccades in complex scenes are made when the odds for the target's presence outside the fovea versus within the fovea are about four.

Cytogenetic characteristics of patients with signs and symptoms of myelodysplastic syndromes in the State of Pará, Brazil

The myelodysplastic syndromes (MDS) are clonal hematopoietic diseases characterized by medullary dysplasia, cytopenias, and frequent evolution to acute myeloid leukemia. In 1982, the French-American-British (FAB) group proposed a classification for the MDS, based on morphological characteristics of peripheral blood and of the bone marrow. Later, cytogenetics proved to be a useful tool for the refinement of prognosis, through the use of the International Prognosis Score System (IPSS), as well as through evidence of clonality. Recently, the World Health Organization (WHO) proposed a new classification for the MDS, based on significant modifications of the FAB proposal, with the inclusion of chromosome analysis. A cytogenetic analysis was made of 17 patients with symptoms of MDS in the State of Para, based on WHO recommendations, and application of the IPSS. Good metaphases were obtained for 13 patients; 12 had a normal karyotype and only one had a clonal abnormality, del(3)(p25). The genes related to neoplastic processes that have been mapped to 3p are: XPC in 3p25.1 and FANCD2 and VHL in 3p25-26. Four patients had classic symptoms of MDS; in the rest the possibility of MDS was excluded or several months of observation before diagnosis were recommended. Among those with MDS, it was not possible to apply IPSS and WHO recommendations, because fundamental data were lacking, specifically the medullary blast and ring sideroblast counts. We advocate the implementation of routine cytogenetic analyses for the study of MDS, especially in patients with moderate hematopoietic dysplasia

Localization of the plane of regard in space.

When we fixate an object in space, the rotation centers of the eyes, together with the object, define a plane of regard. People perceive the elevation of objects relative to this plane accurately, irrespective of eye or head orientation (Poljac et al. (2004) Vision Res, in press). Yet, to create a correct representation of objects in space, the orientation of the plane of regard in space is required. Subjects pointed along an eccentric vertical line on a touch screen to the location where their plane of regard intersected the touch screen positioned on their right. The distance of the vertical line to the subject's eyes varied from 10 to 40 cm. Subjects were sitting upright and fixating one of the nine randomly presented directions ranging from 20 degrees left and down to 20 degrees right and up relative to their straight ahead. The eccentricity of fixations relative to the pointing location varied by up to 40 degrees . Subjects underestimated the elevation of their plane of regard (on average by 3.69 cm, SD=1.44 cm), regardless of the fixation direction or pointing distance. However, when the targets were shown on a display mounted in a table, to provide support of the subject's hand throughout the trial, subjects pointed accurately (average error 0.3 cm, SD=0.8 cm). In addition, head tilt 20 degrees to the left or right did not cause any change in accuracy. The bias observed in the first task could be caused by maintained tonus in arm muscles when the arm is raised, that might interfere with the transformation from visual to motor signals needed to perform the pointing movement. We conclude that the plane of regard is correctly localized in space. This may be a good starting point for representing objects in head-centric coordinates.

Perceptual compensation for eye torsion.

To correctly perceive visual directions relative to the head, one needs to compensate for the eye's orientation in the head. In this study we focus on compensation for the eye's torsion regarding objects that contain the line of sight and objects that do not pass through the fixation point. Subjects judged the location of flashed probe points relative to their binocular plane of regard, the mid-sagittal or the transverse plane of the head, while fixating straight ahead, right upward, or right downward at 30 cm distance, to evoke eye torsion according to Listing's law. In addition, we investigated the effects of head-tilt and monocular versus binocular viewing. Flashed probe points were correctly localized in the plane of regard irrespective of eccentric viewing, head-tilt, and monocular or binocular vision in nearly all subjects and conditions. Thus, eye torsion that varied by +/-9 degrees across these different conditions was in general compensated for. However, the position of probes relative to the midsagittal or the transverse plane, both true head-fixed planes, was misjudged. We conclude that judgment of the orientation of the plane of regard, a plane that contains the line of sight, is veridical, indicating accurate compensation for actual eye torsion. However, when judgment has to be made of a head-fixed plane that is offset with respect to the line of sight, eye torsion that accompanies that eye orientation appears not to be taken into account correctly.

Representation of heading direction in far and near head space.

Manipulation of objects around the head requires an accurate and stable internal representation of their locations in space, also during movements such as that of the eye or head. For far space, the representation of visual stimuli for goal-directed arm movements relies on retinal updating, if eye movements are involved. Recent neurophysiological studies led us to infer that a transformation of visual space from retinocentric to a head-centric representation may be involved for visual objects in close proximity to the head. The first aim of this study was to investigate if there is indeed such a representation for remembered visual targets of goal-directed arm movements. Participants had to point toward an initially foveated central target after an intervening saccade. Participants made errors that reflect a bias in the visuomotor transformation that depends on eye displacement rather than any head-centred variable. The second issue addressed was if pointing toward the centre of a wide-field expanding motion pattern involves a retinal updating mechanism or a transformation to a head-centric map and if that process is distance dependent. The same pattern of pointing errors in relation to gaze displacement was found independent of depth. We conclude that for goal-directed arm movements, representation of the remembered visual targets is updated in a retinal frame, a mechanism that is actively used regardless of target distance, stimulus characteristics or the requirements of the task.

Different visual search strategies in stationary and moving radial patterns.

This study compared visual search strategies in patterns of radially moving dots (simulating self-motion) to those used in matched stationary displays (radial patterns of lines). To control for differences in target visibility, 75% detection thresholds for deviating motion direction and line orientation, respectively, were determined as a function of eccentricity in Experiment 1. These individual thresholds were used to study saccadic parameters in Experiment 2, when subjects searched for targets in the stationary and moving patterns. Despite similarities in search performance, visual search in moving radial patterns was characterised by fewer saccades, longer initial fixation times, and shorter saccadic amplitudes after the initial saccade than during search in a matched stationary radial pattern. These results suggest that detection performance alone cannot explain saccadic search behaviour, and that different search strategies may be used in moving compared to stable environments.

The timing of sequences of saccades in visual search.

According to the LATER model (linear approach to thresholds with ergodic rate), the latency of a single saccade in response to target appearance can be understood as a decision process, which is subject to (i) variations in the rate of (visual) information processing; and (ii) the threshold for the decision. We tested whether the LATER model can also be applied to the sequences of saccades in a multiple fixation search, during which latencies of second and subsequent saccades are typically shorter than that of the initial saccade. We found that the distributions of the reciprocal latencies for later saccades, unlike those of the first saccade, are highly asymmetrical, much like a gamma distribution. This suggests that the normal distribution of the rate r, which the LATER model assumes, is not appropriate to describe the rate distributions of subsequent saccades in a scanning sequence. By contrast, the gamma distribution is also appropriate to describe the distribution of reciprocal latencies for the first saccade. The change of the gamma distribution parameters as a function of the ordinal number of the saccade suggests a lowering of the threshold for second and later saccades, as well as a reduction in the number of target elements analysed.

Heading and path percepts from visual flow and eye pursuit signals.

The percept of self-motion through the environment is supported by visual motion signals and eye movement signals. The interaction between these signals by decoupling of the eye movement and the pattern of retinal motion during brief simulated ego-movement on straight or circular trajectories was studied. A new response method enabled subjects to report perceived destination and perceived curvature of their future path simultaneously. Various combinations of simulated gaze rotation in the retinal flow and eye pursuit were investigated. Simulated gaze rotation ranged from consistent and larger than, to opponent and larger than eye pursuit. It was found that the perceived destination shifts non-linearly with the mismatch between simulated gaze rotation and eye pursuit. The non-linearity is also revealed in the perceived tangent heading direction and perceived path curvature, although to different extent in different subjects. For the same retinal flow, eye pursuit that is consistent with the simulated gaze rotation reduces heading error and the perceived path straightens out. In contrast, perceived path and/or heading do not become more curved or more biased in the direction opposite to pursuit when the eye -in-head rotation is opposite to the simulated gaze rotation. These observations point to modulation of the effect of the extra-retinal pursuit signal by the visual evidence for eye rotation. In a second experiment, one presented to a stationary eye the sum of a component of simulated gaze rotation and radial flow. It was found that the bi-circular flow component, that characterizes the change in pattern of flow directions by the gaze rotation, induces a shift of perceived heading without appreciable perceived path curvature. Conversely, the complementary component of simulated gaze rotation (bi-radial flow) evokes a percept of motion on a curved path with a small tangent heading error. It was suggested that bi-circular and bi-radial flow components contribute primarily to percepts of heading and path curvature, respectively.

Distinguishing subregions of the human MT+ complex using visual fields and pursuit eye movements.

In humans, functional imaging studies have demonstrated a homologue of the macaque motion complex, MT+ [suggested to contain both middle temporal (MT) and medial superior temporal (MST)], in the ascending limb of the inferior temporal sulcus. In the macaque monkey, motion-sensitive areas MT and MST are adjacent in the superior temporal sulcus. Electrophysiological research has demonstrated that while MT receptive fields primarily encode the contralateral visual field, MST dorsal (MSTd) receptive fields extend well into the ipsilateral visual field. Additionally, macaque MST has been shown to receive extraretinal smooth-pursuit eye-movement signals, whereas MT does not. We used functional magnetic resonance imaging (fMRI) and the neural properties that had been observed in monkeys to distinguish putative human areas MT from MST. Optic flow stimuli placed in the full field, or contralateral field only, produced a large cluster of functional activation in our subjects consistent with previous reports of human area MT+. Ipsilateral optic flow stimuli limited to the peripheral retina produced activation only in an anterior subsection of the MT+ complex, likely corresponding to putative MSTd. During visual pursuit of a single target, a large portion of the MT+ complex was activated. However, during nonvisual pursuit, only the anterolateral portion of the MT+ complex was activated. This subsection of the MT+ cluster could correspond to putative MSTl (lateral). In summary, we observed three distinct subregions of the human MT+ complex that were arranged in a manner similar to that seen in the monkey.

Pursuit affects precision of perceived heading for small viewing apertures.

We investigated the interaction between extra-retinal rotation signals and retinal motion signals in heading perception during pursuit eye movement. For limited viewing aperture, the variability in perceived heading strongly depends on the pattern of motion directions. Heading towards a point outside the aperture generates nearly parallel aperture flow. This results in lower precision of perceived heading than heading that renders the radial pattern of flow visible. We ask if the precision is limited by the pattern of flow visible on the retina or that on the screen. During fixation, the two patterns are identical. They are decoupled during pursuit, since pursuit changes radial flow within the aperture on the screen into nearly parallel flow on the retina, and vice versa. The extra-retinal signal is known to reduce systematic errors in the direction of pursuit, thus compensating for the rotational flow during pursuit. We now ask if the extra-retinal signal also affects the precision of heading percepts. It might if at the spatial integration stage the rotational flow has been subtracted out already. A compensation beyond the integration stage, however, cannot undo the change in retinal motion directions so that an effect of pursuit on precision cannot be avoided. We measured the variable and systematic errors in perceived heading during fixation and pursuit for a frontal plane approach, while varying duration, dot lifetime and aperture size. We found precision is effected by pursuit as much as predicted from the pattern of retinal flow, while compensation is significantly greater than zero. This means that the interaction between the extra-retinal signal and visual motion signals takes place after spatial integration of local motion signals. Furthermore, compensation increased significantly with longer duration (0.5-3.0 s), but not with larger aperture size (10-50 degrees ). A larger aperture size did increase the eccentricity of perceived heading.