The cortical visual area, V4, has been considered to code contours that contribute to the intermediate-level representation of objects. The neural responses to the complex contour features intrinsic to natural contours are expected to clarify the essence of the representation. To approach the cortical coding of natural contours, we investigated the simultaneous coding of multiple contour features in monkey (Macaca fuscata) V4 neurons and their population-level representation. A substantial number of neurons showed significant tuning for two or more features such as curvature and closure, indicating that a substantial number of V4 neurons simultaneously code multiple contour features. A large portion of the neurons responded vigorously to acutely curved contours that surrounded the center of classical receptive field, suggesting that V4 neurons tend to code prominent features of object contours. The analysis of mutual information (MI) between the neural responses and each contour feature showed that most neurons exhibited similar magnitudes for each type of MI, indicating that many neurons showing the responses depended on multiple contour features. We next examined the population-level representation by using multidimensional scaling analysis. The neural preferences to the multiple contour features and that to natural stimuli compared with silhouette stimuli increased along with the primary and secondary axes, respectively, indicating the contribution of the multiple contour features and surface textures in the population responses. Our analyses suggested that V4 neurons simultaneously code multiple contour features in natural images and represent contour and surface properties in population.
Keywords: closure; electrophysiology; multiple tuning; mutual information; natural images; simultaneous coding.
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