Gradients of binocular disparity across the visual field provide a potent cue to the three-dimensional (3-D) orientation of surfaces in a scene. Neurons selective for 3-D surface orientation defined by disparity gradients have recently been described in parietal cortex, but little is known about where and how this selectivity arises within the visual pathways. Because the middle temporal area (MT) has previously been implicated in depth perception, we tested whether MT neurons could signal the 3-D orientation (as parameterized by tilt and slant) of planar surfaces that were depicted by random-dot stereograms containing a linear gradient of horizontal disparities. We find that many MT neurons are tuned for 3-D surface orientation, and that tilt and slant generally have independent effects on MT responses. This separable coding of tilt and slant is reminiscent of the joint coding of variables in other areas (e.g., orientation and spatial frequency in V1). We show that tilt tuning remains unchanged when all coherent motion is removed from the visual stimuli, indicating that tilt selectivity is not a byproduct of 3-D velocity coding. Moreover, tilt tuning is typically insensitive to changes in the mean disparity (depth) of gradient stimuli, indicating that tilt tuning cannot be explained by conventional tuning for frontoparallel disparities. Finally, we explore the receptive field mechanisms underlying selectivity for 3-D surface orientation, and we show that tilt tuning arises through heterogeneous disparity tuning within the receptive fields of MT neurons. Our findings show that MT neurons carry high-level signals about 3-D surface structure, in addition to coding retinal image velocities.

Sanada TM, Nguyenkim JD, DeAngelis GC. Representation of 3-D surface orientation by velocity and disparity gradient cues in area MT. J Neurophysiol 107: 2109 -2122, 2012. First published January 4, 2012; doi:10.1152/jn.00578.2011.-Neural coding of the threedimensional (3-D) orientation of planar surface patches may be an important intermediate step in constructing representations of complex 3-D surface structure. Spatial gradients of binocular disparity, image velocity, and texture provide potent cues to the 3-D orientation (tilt and slant) of planar surfaces. Previous studies have described neurons in both dorsal and ventral stream areas that are selective for surface tilt based on one or more of these gradient cues. However, relatively little is known about whether single neurons provide consistent information about surface orientation from multiple gradient cues. Moreover, it is unclear how neural responses to combinations of surface orientation cues are related to responses to the individual cues. We measured responses of middle temporal (MT) neurons to random dot stimuli that simulated planar surfaces at a variety of tilts and slants. Four cue conditions were tested: disparity, velocity, and texture gradients alone, as well as all three gradient cues combined. Many neurons showed robust tuning for surface tilt based on disparity and velocity gradients, with relatively little selectivity for texture gradients. Some neurons showed consistent tilt preferences for disparity and velocity cues, whereas others showed large discrepancies. Responses to the combined stimulus were generally well described as a weighted linear sum of responses to the individual cues, even when disparity and velocity preferences were discrepant. These findings suggest that area MT contains a rudimentary representation of 3-D surface orientation based on multiple cues, with single neurons implementing a simple cue integration rule. depth; slant; surface; tilt; visual cortex; middle temporal area THE VISUAL SYSTEM RECONSTRUCTS three-dimensional (3-D) scene structure from images projected onto the two retinas. Many cues, including binocular disparity, relative motion, texture, shading, and perspective, are used to perceive 3-D structure. Most complex surfaces can be approximated by combinations of locally planar surfaces. Thus understanding how planar surfaces are coded in visual cortex may help reveal how complex surface representations are constructed. The 3-D orientation of a plane (tilt and slant) can be specified by gradients of binocular disparity, motion (velocity), or texture. Human perception of 3-D surface orientation from these cues has been well studied, and the findings are often well explained by Bayesian models (Girshick and Banks 2009;Hillis et al. 2004;Jacobs 1999;Knill 2007;Knill and Saunders 2003).Physiological studies in macaques have identified neurons that signal the 3-D orientation of planar surfaces. In the ventral stream, 3-D orientation tuning has been reported in area V4 for disparity gradients (Hegde and Van Ess...