Coding of shape from shading in area V4 of the macaque monkey

BACKGROUND: The shading of an object provides an important cue for recognition, especially for determining its 3D shape. However, neuronal mechanisms that allow the recovery of 3D shape from shading are poorly understood. The aim of our study was to determine the neuronal basis of 3D shape from shading coding in area V4 of the awake macaque monkey. RESULTS: We recorded the responses of V4 cells to stimuli presented parafoveally while the monkeys fixated a central spot. We used a set of stimuli made of 8 different 3D shapes illuminated from 4 directions (from above, the left, the right and below) and different 2D controls for each stimulus. The results show that V4 neurons present a broad selectivity to 3D shape and illumination direction, but without a preference for a unique illumination direction. However, 3D shape and illumination direction selectivities are correlated suggesting that V4 neurons can use the direction of illumination present in complex patterns of shading present on the surface of objects. In addition, a vast majority of V4 neurons (78%) have statistically different responses to the 3D and 2D versions of the stimuli, while responses to 3D are not systematically stronger than those to 2D controls. However, a hierarchical cluster analysis showed that the different classes of stimuli (3D, 2D controls) are clustered in the V4 cells response space suggesting a coding of 3D stimuli based on the population response. The different illumination directions also tend to be clustered in this space. CONCLUSION: Together, these results show that area V4 participates, at the population level, in the coding of complex shape from the shading patterns coming from the illumination of the surface of corrugated objects. Hence V4 provides important information for one of the steps of cortical processing of the 3D aspect of objects in natural light environment.