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Biomedical subjects

A V van den Berg

Publications and source records attributed to A V van den Berg.

At least 19 recordsLinked to original sources

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.

Animals↗

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.

Attention↗

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.

Analysis of Variance↗

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.

Arm↗

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.

Eye Movements↗

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.

Discrimination, Psychological↗

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.

Arm↗

Smooth eye movements and spatial localisation.

We asked subjects to align a target that flashed as their eyes rotated to the right in pursuit of a moving ring, with a target that flashed as their eyes rotated to the left in pursuit of the ring. Subjects systematically mislocalised the targets in the direction of pursuit. When the ring and flashes were the only structures that were visible, the alignment error was about 4 cm, corresponding to a timing error of about 100 ms. The timing error was independent of the position along the ring's path, but did depend to some extent on pursuit velocity. Adding a textured background reduced the mislocalisation considerably, presumably because it enabled subjects to localise the targets relative to the surrounding. There was almost no mislocalisation if the subject was not pursuing the ring. It is suggested that the mislocalisation arises because incoming retinal signals are combined directly with outgoing oculo-motor commands, with no attempt to account for any of the involved neuronal and muscular delays.

Computer Graphics↗

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.

Humans↗

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.

Discrimination, Psychological↗

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.

Animals↗

Perceived heading during simulated torsional eye movements.

Observer translation through the environment can be accompanied by rotation of the eye about any axis. For rotation about the vertical axis (horizontal rotation) during translation in the horizontal plane, it is known that the absence of depth in the scene and an extra retinal signal leads to a systematic error in the observer's perceived direction of heading. This heading error is related in magnitude and direction to the shift of the centre of retinal flow (CF) that occurs because of the rotation. Rotation about any axis that deviates from the heading direction results in a CF shift. So far, however, the effect of rotation about the line of sight (torsion) on perceived heading has not been investigated. We simulated observer translation towards a wall or cloud, while simultaneously simulating eye rotation about the vertical axis, the torsional axis or combinations thereof. We find only small systematic effects of torsion on the set of 2D perceived headings, regardless of the simulated horizontal rotation. In proportion to the CF shift, the systematic errors are significantly smaller for pure torsion than for pure horizontal rotation. In contrast to errors caused by horizontal rotation, the torsional errors are hardly reduced by addition of depth to the scene. We suggest the difference in behaviour reflects the difference in symmetry of the field of view relative to the axis of rotation: the higher symmetry in the case of torsion may allow for a more accurate estimation of the rotational flow. Moreover, we report a new phenomenon. Simulated horizontal rotation during simulated wall approach increases the heading-dependency of errors, causing a larger compression of perceived heading in the horizontal direction than in the vertical direction.

Eye Movements↗

The mechanism of interaction between visual flow and eye velocity signals for heading perception.

A translating eye receives a radial pattern of motion that is centered on the direction of heading. If the eye is rotating and translating, visual and extraretinal signals help to cancel the rotation and to perceive heading correctly. This involves (1) an interaction between visual and eye movement signals and (2) a motion template stage that analyzes the pattern of visual motion. Early interaction leads to motion templates that integrate head-centered motion signals in the visual field. Integration of retinal motion signals leads to late interaction. Here, we show that retinal flow limits precision of heading. This result argues against an early, vector subtraction type of interaction, but is consistent with a late, gain field type of interaction with eye velocity signals and neurophysiological findings in area MST of the monkey.

Eye Movements↗

Visually evoked cyclovergence and extended listing's law.

Cyclovergence is a simultaneously occurring cyclorotation of the two eyes in opposite directions. Cyclovergence can be elicited visually by opposite cyclorotation of the two eyes' images. It also can occur in conjunction with horizontal vergence and vertical version in a stereotyped manner as described by the extended Listing's law (or L2). We manipulated L2-related and visually evoked cyclovergence independently, using stereoscopic images of three-dimensional (3D) scenes. During pursuit in the midsagittal plane, cyclovergence followed L2. The amount of L2-related cyclovergence during pursuit varied between subjects. Each pursuit trial was repeated three times. Two of the three trials had additional image rotation to visually evoke cyclovergence. We could separate the L2-related and visual components of cyclovergence by subtraction of the cyclovergence response in matched trials that differed only in the image rotation that was applied during pursuit. This indicates that visual and L2-related contributions to cyclovergence add linearly, suggesting the presence of two independent systems. Visually evoked cyclovergence gains were characteristic for a given subject, little affected by visual stimulus parameters, and usually low (0.1-0.5) when a static target was fixated. Gain and phase lag of the visually evoked cyclovergence during vertical pursuit was comparable with that during fixation of a static target. The binocular orientations are in better agreement to orientations predicted by L2 then would be predicted by nulling of the cyclodisparities. On the basis of our results, we suggest that visually driven and L2-related cyclovergence are independent of each other and superimpose linearly.

Adult↗

Visual search of heading direction.

When we move along we frequently look around. How quickly and accurately can we gaze in the direction of heading? We studied the temporal aspects of heading perception in expanding and contracting patterns simulating self-motion. Center of flow (CF) eccentricity was 15 degrees. Subjects had to indicate the CF by making a saccade to it. A temporal constraint on the response time was introduced, because stimuli were presented briefly (1 s). On average, subjects needed two saccades to find the CF. Initial saccades covered about 50-60% of the distance between the fixation point and the CF. Subjects underestimated the eccentricity of the CF. The systematic radial error ranged from -2.4 degrees to -4.9 degrees. The systematic tangential error was small (about 0.5 degree). The variable radial error ranged from 2.7 degrees to 4.6 degrees. We found a relation between saccade onset time and saccade endpoint error. Saccade endpoint error decreased with increasing saccade onset time, suggesting that saccades were often fired before the heading processing had been completed. From the saccade onset times, saccade endpoint errors and an estimate for the saccadic dead time (interval prior to the saccade during which modification is impossible 70 ms), we estimated the heading processing time (HPT 0.43 s). In three out of four subjects, HPT was longer for trials simulating backward movement than for trials simulating forward movement. For each saccade we determined whether it reduced the distance error. The second saccade reduced the error more effectively per time unit than the initial saccade. On the basis of this finding, we suggest that visual processing that occurs during the saccadic dead time of the first saccade is used in the preparation of the second saccade.

Adult↗

Predicting the present direction of heading.

Humans perceive heading accurately when they rotate their eyes. This is remarkable, because (1) the pursuit eye movement makes the retinal flow more complicated; and (2) the eye rotation causes a continuous change of the heading direction on the retina. The first problem prevents a simple association of the centre of flow on the retina with the heading direction. To solve it, the brain needs to take into account the flow associated with the eye's rotation. But even if this is done correctly, the resulting estimate of the heading is retino-centric and changing over time. Thus, the processing time to retrieve the heading from the flow field will cause a lag with respect to the actual heading direction. We investigated the latency for heading perception. We presented step wise changes of the centre of expanding flow to stationary and moving eyes. This mimics the movement of the heading direction across the retina, but avoids the complicating effects of rotational flow. For a stationary eye, we found a bias in perceived heading that corresponds to a latency of 300 ms or more. Yet, errors in heading perception are marginal normally, because we found an opposite bias for the moving eye, which counters the errors due to latency and a changing retino-centric heading direction. This suggests that the current heading direction is predicted from the extra-retinal signal and the delayed visual signals.

Eye Movements↗

Local and global factors affecting the coherent motion of gratings presented in multiple apertures.

Using stimuli composed of two independent gratings viewed through multiple apertures, we investigate a number of parameters affecting the integration of locally ambiguous motions into globally coherent motion. In four experiments, we varied local factors (grating spatial frequency, speed, contrast, duty cycle, orientation) and global factors (degree of similarity and common fate between the gratings, and symmetry in the configuration of the grating pattern) and examined their effects on global motion coherence. Our results, confirming accounts offered by previous investigators, indicate that local competition between motion signals generated by contours (ambiguous) and their line terminations (unambiguous) is important in determining global motion coherence in multiple-aperture stimuli. Our results also indicate that global factors can affect perceived coherence independently of local motion signals, suggesting the involvement of higher-level motion areas and a role for non-motion processes such as those involved in pattern and form perception. Comparing motion coherence with other two-dimensional (2-D) stimuli (plaids) shows that 2-D multiple-aperture stimuli are not analogous and that coherence models derived from plaid stimuli do not account for the data.

Humans↗