Huseyin Boyaci

The representation of perceived angular size in human primary visual cortex

Two objects that project the same visual angle on the retina can appear to occupy very different proportions of the visual field if they are perceived to be at different distances. What happens to the retinotopic map in primary visual cortex (V1) during the perception of these size illusions? Here we show, using functional magnetic resonance imaging (fMRI), that the retinotopic representation of an object changes in accordance with its perceived angular size. A distant object that appears to occupy a larger portion of the visual field activates a larger area in V1 than an object of equal angular size that is perceived to be closer and smaller. These results demonstrate that the retinal size of an object and the depth information in a scene are combined early in the human visual system.

Attention-Dependent Representation of a Size Illusion in Human V1

One of the most fundamental properties of human primary visual cortex (V1) is its retinotopic organization, which makes it an ideal candidate for encoding spatial properties, such as size, of objects. However, three-dimensional (3D) contextual information can lead to size illusions that are reflected in the spatial pattern of activity in V1 [1]. A critical question is how complex 3D contextual information can influence spatial activity patterns in V1. Here, we assessed whether changes in the spatial distribution of activity in V1 depend on the focus of attention, which would be suggestive of feedback of 3D contextual information from higher visual areas. We presented two 3D rings at close and far apparent depths in a 3D scene. When subjects fixated its center, the far ring appeared to be larger and occupy a more eccentric portion of the visual field, relative to the close ring. Using functional magnetic resonance imaging, we found that the spatial distribution of V1 activity induced by the far ring was also shifted toward a more eccentric representation of the visual field, whereas that induced by the close ring was shifted toward the foveal representation, consistent with their perceptual appearances. This effect was significantly reduced when the focus of spatial attention was narrowed with a demanding central fixation task. We reason that focusing attention on the fixation task resulted in reduced activity in--and therefore reduced feedback from--higher visual areas that process the 3D depth cues.

Recessive LAMC3 mutations cause malformations of occipital cortical development

The biological basis for regional and inter-species differences in cerebral cortical morphology is poorly understood. We focused on consanguineous Turkish families with a single affected member with complex bilateral occipital cortical gyration abnormalities. By using whole-exome sequencing, we initially identified a homozygous 2-bp deletion in LAMC3, the laminin γ3 gene, leading to an immediate premature termination codon. In two other affected individuals with nearly identical phenotypes, we identified a homozygous nonsense mutation and a compound heterozygous mutation. In human but not mouse fetal brain, LAMC3 is enriched in postmitotic cortical plate neurons, localizing primarily to the somatodendritic compartment. LAMC3 expression peaks between late gestation and late infancy, paralleling the expression of molecules that are important in dendritogenesis and synapse formation. The discovery of the molecular basis of this unusual occipital malformation furthers our understanding of the complex biology underlying the formation of cortical gyrations.

Border ownership selectivity in human early visual cortex and its modulation by attention

Natural images are usually cluttered because objects occlude one another. A critical aspect of recognizing these visual objects is to identify the borders between image regions that belong to different objects. However, the neural coding of border ownership in human visual cortex is largely unknown. In this study, we designed two simple but compelling stimuli in which a slight change of contextual information could induce a dramatic change of border ownership. Using functional MRI adaptation, we found that border ownership selectivity in V2 was robust and reliable across subjects, and it was largely dependent on attention. Our study provides the first human evidence that V2 is a critical area for the processing of border ownership and that this processing depends on the modulation from higher-level cortical areas.

Attention modulates neuronal correlates of interhemispheric integration and global motion perception

In early retinotopic areas of the human visual system, information from the left and right visual hemifields (VHFs) is processed contralaterally in two hemispheres. Despite this segregation, we have the perceptual experience of a unified, coherent, and uninterrupted single visual field. How exactly the visual system integrates information from the two VHFs and achieves this perceptual experience still remains largely unknown. In this study using fMRI, we explored candidate areas that are involved in interhemispheric integration and the perceptual experience of a unified, global motion across VHFs. Stimuli were two-dimensional, computer-generated objects with parts in both VHFs. The retinal image in the left VHF always remained stationary, but in the experimental condition, it appeared to have local motion because of the perceived global motion of the object. This perceptual effect could be weakened by directing the attention away from the global motion through a demanding fixation task. Results show that lateral occipital areas, including the medial temporal complex, play an important role in the process of perceptual experience of a unified global motion across VHFs. In early areas, including the lateral geniculate nucleus and V1, we observed correlates of this perceptual experience only when attention is not directed away from the object. These findings reveal effects of attention on interhemispheric integration in motion perception and imply that both the bilateral activity of higher-tier visual areas and feedback mechanisms leading to bilateral activity of early areas play roles in the perceptual experience of a unified visual field.

Human observers compensate for secondary illumination originating in nearby chromatic surfaces

In complex scenes, the light absorbed and re-emitted by one surface can serve as a source of illumination for a second. We examine whether observers systematically discount this secondary illumination when estimating surface color. We asked six naive observers to make achromatic settings of a small test patch adjacent to a brightly colored orange cube in rendered scenes. The orientation of the test patch with respect to the cube was varied from trial to trial, altering the amount of secondary illumination reaching the test patch. Observers systematically took orientation into account in making their settings, discounting the added secondary illumination more at orientations where it was more intense. Overall, they tended to under-compensate for the added secondary illumination.

Cues to an Equivalent Lighting Model

We investigate how human observers make use of three candidate cues in their lightness judgments. Each cue potentially provides information about the spatial distribution of light sources in complex, rendered 3D scenes. The illumination (lighting model) of each scene consisted of a punctate light source combined with a diffuse light source. The cues were (1) cast shadows, (2) surface shading, and (3) specular highlights. Observers were asked to judge the albedo of a matte grayscale test patch that varied in orientation with respect to the punctate light source. We tested their performance in scenes containing only one type of cue and in scenes where all three cue types were present. From the results, we deduced how accurately they had estimated the spatial distribution of light sources in each scene given the cues available. In Experiment 1, we established that all of the individual cues were used in isolation. We showed that the highlight and cast shadow cues in isolation were used by more than half of the observers. We could reject the hypothesis that the observers did not make use of the shading cue for only one observer. In Experiment 2, we showed that the observers combined information from multiple cues when all three cues were presented together.

Testing limits on matte surface color perception in three-dimensional scenes with complex light fields.

We investigated limits on the human visual systems ability to discount directional variation in complex lights field when estimating Lambertian surface color. Directional variation in the light field was represented in the frequency domain using spherical harmonics. The bidirectional reflectance distribution function of a Lambertian surface acts as a low-pass filter on directional variation in the light field. Consequently, the visual system needs to discount only the low-pass component of the incident light corresponding to the first nine terms of a spherical harmonics expansion [Basri, R., Jacobs, D. (2001). Lambertian reflectance and linear subspaces. In: International Conference on Computer Vision II, pp. 383-390; Ramamoorthi, R., Hanrahan, P., (2001). An efficient representation for irradiance environment maps. SIGGRAPH 01. New York: ACM Press, pp. 497-500] to accurately estimate surface color. We test experimentally whether the visual system discounts directional variation in the light field up to this physical limit. Our results are consistent with the claim that the visual system can compensate for all of the complexity in the light field that affects the appearance of Lambertian surfaces.

Perceived glossiness in high dynamic range scenes

We investigated how spatial pattern, background, and dynamic range affect perceived gloss in brightly lit real scenes. Observers viewed spherical objects against uniform backgrounds. There were three possible objects. Two were black matte spheres with circular matte white dots painted on them (matte-dot spheres). The third sphere was painted glossy black (glossy black sphere). Backgrounds were either black or white matte, and observers saw each of the objects in turn on each background. Scenes were illuminated by an intense collimated source. On each trial, observers matched the apparent albedo of the sphere to an albedo reference scale and its apparent gloss to a gloss reference scale. We found that matte-dot spheres and the black glossy sphere were perceived as glossy on both backgrounds. All spheres were judged to be significantly glossier when in front of the black background. In contrast with previous research using conventional computer displays, we find that background markedly affects perceived gloss. This finding is surprising because darker surfaces are normally perceived as glossier (F. Pellacini, J. A. Ferwerda, & D. P. Greenberg, 2000). We conjecture that there are cues to surface material signaling glossiness present in high dynamic range scenes that are absent or weak in scenes presented using conventional computer displays.

Vision in 3D Environments: Surface color perception and light field estimation in 3D scenes

Previous research on surface color perception has typically used Mondrian stimuli consisting of a small number of matte surface patches in a plane perpendicular to the line of sight. In such scenes, reliable estimation of the color of a surface is a difficult if not impossible computational problem (Maloney, 1999). In three-dimensional scenes consisting of surfaces at many different orientations it is at least in theory possible to estimate surface color. However, the difficulty of the problem increases, in part, because the effective illumination incident on each surface (the light field) now depends on surface orientation and location. We review recent work in multiple laboratories that examines the degree to which the human visual system discounts the light field in judging matte surface lightness and color and how the visual system estimates the flow of light in a scene.