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R von der Heydt

Publications and source records attributed to R von der Heydt.

At least 19 recordsLinked to original sources
PubMed2000

Coding of border ownership in monkey visual cortex.

Areas V1 and V2 of the visual cortex have traditionally been conceived as stages of local feature representations. We investigated whether neural responses carry information about how local features belong to objects. Single-cell activity was recorded in areas V1, V2, and V4 of awake behaving monkeys. Displays were used in which the same local feature (contrast edge or line) could be presented as part of different figures. For example, the same light-dark edge could be the left side of a dark square or the right side of a light square. Each display was also presented with reversed contrast. We found significant modulation of responses as a function of the side of the figure in >50% of neurons of V2 and V4 and in 18% of neurons of the top layers of V1. Thus, besides the local contrast border information, neurons were found to encode the side to which the border belongs ("border ownership coding"). A majority of these neurons coded border ownership and the local polarity of luminance-chromaticity contrast. The others were insensitive to contrast polarity. Another 20% of the neurons of V2 and V4, and 48% of top layer V1, coded local contrast polarity, but not border ownership. The border ownership-related response differences emerged soon (<25 msec) after the response onset. In V2 and V4, the differences were found to be nearly independent of figure size up to the limit set by the size of our display (21 degrees ). Displays that differed only far outside the conventional receptive field could produce markedly different responses. When tested with more complex displays in which figure-ground cues were varied, some neurons produced invariant border ownership signals, others failed to signal border ownership for some of the displays, but neurons that reversed signals were rare. The influence of visual stimulation far from the receptive field center indicates mechanisms of global context integration. The short latencies and incomplete cue invariance suggest that the border-ownership effect is generated within the visual cortex rather than projected down from higher levels.

Animals↗
PubMed2000

Representation of stereoscopic edges in monkey visual cortex.

Form perception in random-dot stereograms is based on information that resides in the correlation between the two images, but is not present in either image alone. We have studied the coding of stereoscopic figures in the neural activity of areas V1 and V2 of alert behaving monkeys. While cells in V1 generally responded according to the disparity of the surface at the receptive field, we found cells in area V2 that responded selectively to the figure edges. These cells signaled the location and orientation of contrast borders as well as stereoscopic edges, and were often selective for the direction of the step in depth. We concluded that stereoscopic edges are explicitly represented in area V2.

Animals↗
PubMed1999

Color filling-in under steady fixation: behavioral demonstration in monkeys and humans.

Color filling-in is a phenomenon in which the color of an object appears to be filled-in by the color of the surrounding field. We have studied the question of whether monkeys perceive color filling-in with near-foveal stimuli under steady fixation. Two monkeys were trained to fixate steadily and to attend to a disk, surrounded by an annulus of the complementary color, in parafoveal vision. Using displays in which the color of the disk was gradually changed to that of the annulus, we trained the animals to signal when they perceived a uniform color field. During the experiment, we introduced a small percentage of trials in which the disk color remained constant, and looked for 'filling-in' responses in these trials. Three human subjects were also tested for comparison. All subjects produced 'filling-in' responses with frequencies that were significantly higher for static disks with blurred borders than for moving disks or disks with sharp borders. This indicates that the monkeys' responses reflected perceptual filling-in, rather than random behavior. The time course of filling-in was similar in monkeys and humans. For the blurred static disks, responses occurred first after 3-4 s of fixation, reaching a probability of 0.2-0.8 by the end of 6 s, depending on the subject.

Adult↗
PubMed1993

Functional organization of area V2 in the alert macaque.

We studied the relation between anatomical structure and functional properties of cells in area V2 of the macaque. Visual function was assessed in the alert animal during fixation of gaze. Recording sites were reconstructed with respect to cortical lamination and the cytochrome oxidase pattern. We measured orientation and direction selectivity, end-stopping, sensitivity to binocular disparity and ocular dominance, and determined more complex functions like sensitivity to anomalous contours and lines defined by coherent motion. Orientation selectivity was found in all parts of area V2, with high frequencies in the pale and thick stripes of the cytochrome oxidase pattern, and with lower frequency in the thin stripes. Representations of anomalous contours were found in the pale and thick stripes with similar frequencies, but generally not in the thin stripes, which have been thought to process colour. Lines defined by coherent motion were most frequently represented in the thick stripes; they were less frequent in the pale stripes, and (as with anomalous contours) were not found in the thin stripes. Sensitivity to binocular disparity was found in all types of stripes, but more frequently in the thick stripes, where the exclusively binocular neurons were also concentrated. By contrast, no segregation was found for direction selectivity and end-stopping. All neuronal properties were distributed evenly across cortical laminae. We conclude that mechanisms for figure-ground segregation involve the pale and the thick stripes of the cytochrome oxidase pattern, perhaps with greater emphasis on 'shape from motion' and 'stereoscopic depth' in the thick stripes, while more elementary neuronal properties are distributed almost evenly across the stripe pattern.

Animals↗
PubMed1992

Simulation of neural contour mechanisms: from simple to end-stopped cells.

Early stages of visual form processing were modelled by simulating cortical simple, complex and end-stopped cells. The computation involves (1) convolution of the image with even and odd symmetrical orientation selective filters (S-operators), (2) combination of even and odd filter outputs to a local energy measure (C-operator), (3) "differentiation" of the C-operator maps along the respective orientation (single and double end-stopped operators) and (4) determination of local maxima ("key-points") of the combined end-stopped operator activity. While S- and C-operators are optimised for the representation of 1-D features such as edges and lines, the end-stopped operator responses at the key-points make explicit 2-D signal variations such as line ends, corners and segments of strong curvature. The theoretical need for this complementary representation is discussed. The model was tested on grey-valued images.

Filtration↗
PubMed1992

Periodic-pattern-selective cells in monkey visual cortex.

To study the visual processing of periodic and aperiodic patterns, we have analyzed neuronal responses in areas V1 and V2 of the visual cortex of alert monkeys during behaviorally induced fixation of gaze. Receptive field eccentricities ranged between 0.5 degrees and 4 degrees. We found cells that responded vigorously to gratings, but weakly or not all to bars and edges. In some cells the aperiodic stimuli even reduced the activity below the spontaneous level. The distribution of a bar-grating response index indicated a discrete population of "grating cells" characterized by more than 10-fold superiority of gratings. We estimated that these cells have a frequency of 4% in V1 and 1.6% in V2, and that about 4 million grafting cells of V1 subserve the central 4 degrees of vision. The converse, cells that responded to isolated bars but not to gratings of any periodicity, was also observed. The grating cells of V1 were mostly (23 of 26) found in layers 2, 3, and 4B. They preferred spatial frequencies between 2.6 and 19 cycles/degree (median, 9.3), with tuning widths at half-amplitude between 0.4 and 1.4 octaves (median, 1.0). Their tunings were narrower, and their preferred frequencies higher, than those of other cells on average. Grating cells were also narrowly tuned for orientation. Those of V2 were similarly selective. The responses of grating cells depended critically on the number of cycles of the gratings. With square waves of optimum periodicity responses required a minimum of 2-6 grating cycles and leveled off at 4-14 (median, 7.5). The corresponding receptive field widths were 0.34-2.4 degrees (median, 0.78 degrees) for V1 and 0.72-2.4 degrees (median, 1.4 degrees) for V2. Grating cells typically gave unmodulated responses to drifting gratings, were unselective for direction of motion, and were strongly activated also by stationary gratings. Half of those of V1 were monocular, the others binocular, some showing strong binocular facilitation and disparity sensitivity. Length summation was usually monotonic, but strong end-inhibition was also observed. In contrast to other cells, grating cells were not activated by harmonic components. Spatial-frequency response curves for sine-wave, square-wave, and line gratings were similar. Square-wave gratings of one-third the preferred frequency failed to excite the cells, while the isolated 3f component (f = the fundamental of the square wave) of these gratings evoked strong responses. In spite of the nonlinear features, grating cells had low contrast thresholds.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1991

Subjective contours--bridging the gap between psychophysics and physiology.

Much is known about the initial stages of visual processing up to the striate cortex, but how is visual information represented and handled at subsequent stages? Phenomena of contour, color and movement perception have been used to identify functions of neurons and to reveal functional differences between cortical areas that application of classical receptive-field concepts has not suggested. These differences can be related to theoretical stages of visual processing that provide stability of perception under changing conditions of stimulation.

Animals↗
PubMed1989

Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity.

We have studied the mechanism of contour perception by recording from neurons in the visual cortex of alert rhesus monkeys. In order to assess the relationship between neural signals and perception, we compared the responses to edges and lines with the responses to patterns in which human observers perceive a contour where no line or edge is given (anomalous contour), such as the border between gratings of thin lines offset by half a cycle. With only one exception out of 60, orientation-selective neurons in area V1 did not signal the anomalous contour. Many neurons failed to respond to this stimulus at all, others responded according to the orientation of the grating lines. In area V2, 45 of 103 neurons (44%) signaled the orientation of the anomalous contour. Sixteen did so without signaling the orientation of the inducing lines. Some responded better to anomalous contours than to the optimum bars or edges. Preferred orientations and widths of tuning for anomalous contour and bar or edge were found to be highly correlated, but not identical, in each neuron. Similar to perception, the neuronal responses depended on a minimum number of lines inducing the contour, but not so much on line spacing, and tended to be weaker when the lines were oblique rather than orthogonal to the border. With oblique lines, the orientations signaled were biased towards the orientation orthogonal to the lines, as in the Zöllner illusion. We conclude that contours may be defined first at the level of V2. While the unresponsiveness of neurons in V1 to this type of anomalous contour is in agreement with linear filter predictions, the responses of V2 neurons need to be explained. We assume that they sum the signals of 2 parallel paths, one that defines edges and lines and another that defines anomalous contours by pooling signals from end-stopped receptive fields oriented mainly orthogonal to the contour.

Animals↗
PubMed1989

Mechanisms of contour perception in monkey visual cortex. II. Contours bridging gaps.

We have studied the mechanism of contour perception by recording from neurons in the visual cortex of alert rhesus monkeys. We used stimuli in which human observers perceive anomalous contours: A moving pair of notches in 2 bright rectangles mimicked an overlaying dark bar. For control, the notches were closed by thin lines so that the anomalous contours disappeared or half of the figure was blanked, with a similar effect. Orientation-selective neurons were studied. With the receptive fields centered in the gap, 23 of 72 (32%) neurons tested in area V2 responded to the moving "bar" even though the stimulus spared their response fields, and when the notches were closed, their responses were reduced or abolished. Likewise, when half of the figure was removed, the neurons usually failed to respond. Neurons with receptive fields within 4 degrees of the fovea signaled anomalous contours bridging gaps of 1 degree-3.5 degrees. The anomalous-contour responses were compared quantitatively with response field profiles and length-summation curves and found to exceed the predictions by linear-summation and summation-to-threshold models. Summation models also fail to explain the effect of closing lines which add only negligible amounts of light. In V1, only one of 26 neurons tested showed comparable responses, and only with a narrow gap. The others responded only when the stimulus invaded the response field and did not show the effect of closing lines, or failed to respond at all. The contour responses in V2, the nonadditivity, and the effect of closure can be explained by the previously proposed model (Peterhans et al., 1986), assuming that the corners excite end-stopped fields orthogonal to the contour whose signals are pooled in the contour neurons.

Animals↗
PubMed1984

Illusory contours and cortical neuron responses.

Figures in which human observers perceive "illusory contours" were found to evoke responses in cells of area 18 in the visual cortex of alert monkeys. The cells responded as if the contours were formed by real lines or edges. Modifications that weakened the perception of contours also reduced the neuronal responses. In contrast, cells in area 17 were apparently unable to "see" these contours.

Animals↗
PubMed1983

Plasticity in the binocular correspondence of striate cortical receptive fields in kittens.

The influence of visual experience on the correspondence in position and orientation of receptive fields in the two eyes of cortical neurones was studied. Kittens were reared viewing the environment through lenses that magnified the image by 9% in one direction (meridional size lenses) with axes of magnification oriented 45 degrees left and right of vertical for the two eyes. The unequal deformations in the two eyes produced gradients of position disparity and systematic variation of orientation disparity which could not be influenced by eye movements. Two types of arrangement of the lenses, producing opposite disparities, were used; each was worn by two kittens. The receptive fields of cortical neurones were studied in the four kittens aged 3-4 months. In the binocular cells, the positions of the response fields were plotted, and the preferred orientations determined, using automatic stimulus variation, quantitative analysis, and eye-drift correction. By means of regression analysis, the degree of 'interocular deformation' was assessed; a coefficient D was derived from the positions, an angle beta from the orientations. D specified the position incongruity as a fraction of retinal eccentricity, beta the difference between the orientation incongruities of cells with near-vertical and near-horizontal receptive fields. Both D and beta were found to be of opposite signs in the two groups of kittens, as predicted by the optical effects of the lenses. The difference in D between the groups was 0.197 (predicted: 0.172); the difference in beta was 17.0 degrees (predicted: 18.9 degrees). Thus, the difference in visual environment had been completely compensated by adjustments in the positions as well as the orientations of the receptive fields. Since D and beta are independent of the alignment of the eyes, the differences between the groups reflected different functional connexions at the cortical level. Possible advantages of plasticity for the development of binocular vision are discussed. It is argued that the plasticity demonstrated here reflects a more general property of cortical receptive fields.

Animals↗
PubMed1978

Movement aftereffects in the visual cortex.

Transient effects of prolonged stimulation with moving gratings were studied in single units of the cat striate cortex. In most units, differential adaptation aftereffects of opposite directions of motion could be demonstrated. These results correlate with the psychophysical movement aftereffects.

Action Potentials↗