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Journal of Neuroscience, Vol 9, 1731-1748, Copyright © 1989 by Society for Neuroscience
Mechanisms of contour perception in monkey visual cortex. I. Lines of pattern discontinuity
R von der Heydt and E Peterhans
Department of Neurology, University Hospital Zurich, Switzerland.
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 Zollner 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.
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