Neural correlates of dynamic lightness induction.

The lightness of a surface depends not only on the amount of light reflected off, it but also on the context in which it is embedded. Despite a long history of research, neural correlates of context-dependent lightness perception remain a topic of ongoing debate. Here, we seek to expand on the existing literature by measuring functional magnetic resonance imaging (fMRI) responses to lightness variations induced by the context. During the fMRI experiment, we presented 10 participants with a dynamic stimulus in which either the luminance of a disk or its surround is modulated at four different frequencies ranging from 1 to 8 Hz. Behaviorally, when the surround luminance is modulated at low frequencies, participants perceive an illusory change in the lightness of the disk (lightness induction). In contrast, they perceive little or no induction at higher frequencies. Using this frequency dependence and controlling for long-range responses to border contrast and luminance changes, we found that activity in the primary visual cortex (V1) correlates with lightness induction, providing further evidence for the involvement of V1 in the processing of context-dependent lightness.

Dynamic dot displays reveal material motion network in the human brain.

There is growing research interest in the neural mechanisms underlying the recognition of material categories and properties. This research field, however, is relatively more recent and limited compared to investigations of the neural mechanisms underlying object and scene category recognition. Motion is particularly important for the perception of non-rigid materials, but the neural basis of non-rigid material motion remains unexplored. Using fMRI, we investigated which brain regions respond preferentially to material motion versus other types of motion. We introduce a new database of stimuli - dynamic dot materials - that are animations of moving dots that induce vivid percepts of various materials in motion, e.g. flapping cloth, liquid waves, wobbling jelly. Control stimuli were scrambled versions of these same animations and rigid three-dimensional rotating dots. Results showed that isolating material motion properties with dynamic dots (in contrast with other kinds of motion) activates a network of cortical regions in both ventral and dorsal visual pathways, including areas normally associated with the processing of surface properties and shape, and extending to somatosensory and premotor cortices. We suggest that such a widespread preference for material motion is due to strong associations between stimulus properties. For example viewing dots moving in a specific pattern not only elicits percepts of material motion; one perceives a flexible, non-rigid shape, identifies the object as a cloth flapping in the wind, infers the object's weight under gravity, and anticipates how it would feel to reach out and touch the material. These results are a first important step in mapping out the cortical architecture and dynamics in material-related motion processing.

Distinct patterns of surround modulation in V1 and hMT.

Modulation of a neuron's responses by the stimuli presented outside of its classical receptive field is ubiquitous in the visual system. This "surround modulation" mechanism is believed to be critical for efficient processing and leads to many well-known perceptual effects. The details of surround modulation, however, are still not fully understood. One of the open questions is related to the differences in surround modulation mechanisms in different cortical areas, and their interactions. Here we study patterns of surround modulation in primary visual cortex (V1) and middle temporal complex (hMT+) utilizing a well-studied effect in motion perception, where human observers' ability to discriminate the drift direction of a grating improves as its size gets bigger if the grating has a low contrast, and deteriorates if it has a high contrast. We first replicated the findings in the literature with a behavioral experiment using small and large (1.67 and 8.05 degrees of visual angle) drifting gratings with either low (2%) or high (99%) contrast presented at the periphery. Next, using functional MRI, we found that in V1 with increasing size cortical responses increased at both contrast levels. Whereas in hMT+ with increasing size cortical responses remained unchanged or decreased at high contrast, and increased at low contrast, reflecting the perceptual effect. We also show that the divisive normalization model successfully predicts these activity patterns, and establishes a link between the behavioral results and hMT+ â€‹activity. We conclude that surround modulation patterns in V1 and hMT+ â€‹are different, and that the size-contrast interaction in motion perception is likely to originate in hMT+.

Homozygous LAMC3 mutation links to structural and functional changes in visual attention networks.

The occipital lobe contains a substantial part of the neural machinery involved in visual perception. Mutations in the LAMC3 gene have recently been shown to cause complex bilateral occipital cortical gyration abnormalities. However, to what extent these structural changes impact visual behavior is not known. We recorded responses for two screening test batteries targeting visual function (Leuven - Perceptual Organization Screening Test, Cortical Vision Screening Test) and measured eye fixation performance in a visual attention experiment from a patient with homozygous LAMC3 gene mutation. Using voxel-based morphometry (VBM) we quantitatively assessed the extent of structural changes brought on by the genetic mutation by comparing mean cortical curvature, cortical thickness, and gray matter volume in 34 cortical areas between patient and an age-, sex-, and education-matched control group. Anatomical connectivity between these cortical areas was investigated by a structural covariance analysis. Visual screening-, and behavioral results revealed that the patient's impairments were predominantly in visuo-spatial attention. Consistent with this, VBM and structural connectivity results revealed significant structural changes in cortical regions subserving attentional functions. We conclude that the LAMC3 gene mutation affects cortical areas beyond the occipital lobe and primarily those visual functions that involve heavily distributed networks - such as visuo-spatial attention.

Integrated assessment of visual perception abnormalities in psychotic disorders and relationship with clinical characteristics.

BACKGROUND: The visual system is recognized as an important site of pathology and dysfunction in schizophrenia. In this study, we evaluated different visual perceptual functions in patients with psychotic disorders using a potentially clinically applicable task battery and assessed their relationship with symptom severity in patients, and with schizotypal features in healthy participants. METHODS: Five different areas of visual functioning were evaluated in patients with schizophrenia and schizoaffective disorder (n = 28) and healthy control subjects (n = 31) using a battery that included visuospatial working memory (VSWM), velocity discrimination (VD), contour integration, visual context processing, and backward masking tasks. RESULTS: The patient group demonstrated significantly lower performance in VD, contour integration, and VSWM tasks. Performance did not differ between the two groups on the visual context processing task and did not differ across levels of interstimulus intervals in the backward masking task. Performances on VSWM, VD, and contour integration tasks were correlated with negative symptom severity but not with other symptom dimensions in the patient group. VSWM and VD performances were also correlated with negative sychizotypal features in healthy controls. CONCLUSION: Taken together, these results demonstrate significant abnormalities in multiple visual processing tasks in patients with psychotic disorders, adding to the literature implicating visual abnormalities in these conditions. Furthermore, our results show that visual processing impairments are associated with the negative symptom dimension in patients as well as healthy individuals.

Homozygosity mapping and targeted genomic sequencing reveal the gene responsible for cerebellar hypoplasia and quadrupedal locomotion in a consanguineous kindred.

The biological basis for the development of the cerebro-cerebellar structures required for posture and gait in humans is poorly understood. We investigated a large consanguineous family from Turkey exhibiting an extremely rare phenotype associated with quadrupedal locomotion, mental retardation, and cerebro-cerebellar hypoplasia, linked to a 7.1-Mb region of homozygosity on chromosome 17p13.1-13.3. Diffusion weighted imaging and fiber tractography of the patients' brains revealed morphological abnormalities in the cerebellum and corpus callosum, in particular atrophy of superior, middle, and inferior peduncles of the cerebellum. Structural magnetic resonance imaging showed additional morphometric abnormalities in several cortical areas, including the corpus callosum, precentral gyrus, and Brodmann areas BA6, BA44, and BA45. Targeted sequencing of the entire homozygous region in three affected individuals and two obligate carriers uncovered a private missense mutation, WDR81 p.P856L, which cosegregated with the condition in the extended family. The mutation lies in a highly conserved region of WDR81, flanked by an N-terminal BEACH domain and C-terminal WD40 beta-propeller domains. WDR81 is predicted to be a transmembrane protein. It is highly expressed in the cerebellum and corpus callosum, in particular in the Purkinje cell layer of the cerebellum. WDR81 represents the third gene, after VLDLR and CA8, implicated in quadrupedal locomotion in humans.

Effect of contiguity and figure-ground organization on the area rule of lightness.

In a simple two-dimensional (2D) display composed of two uniform surfaces with different luminances, the lightness of the darker surface varies as a function of its relative area while its luminance is held constant (Gilchrist & Radonjic, 2009; Li & Gilchrist, 1999). This phenomenon is known as the area rule of lightness, and although it is extensively studied in the literature, the underlying principles are still largely unknown. Here, using computer-generated stimuli, we investigated the effects of contiguity and figure-ground organization on the area rule of lightness. Stimuli were 2D disks composed of radial sectors with high (25 cd/m2) or low (8 cd/m2) luminance. On each trial, observers judged the lightness of the sectors by adjusting the luminance of a matching patch. Four conditions were tested. In the contiguous condition, there were one dark and one light sector; in the noncontiguous condition, both the light and dark surfaces were split into four equal radial sectors. Figure and ground conditions were generated by adding small contextual elements to the stimulus. We found that the area rule applied under all conditions; however, the functional form of the effect showed marked differences across conditions. Taken together, our results show that both high-level (e.g., perceptual grouping, figure-ground organization) and low-level (e.g., spatial-summation) mechanisms play a role in the area rule of lightness.

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.

Unmet expectations about material properties delay perceptual decisions.

Based on our expectations about material properties, we can implicitly predict an object's future states, e.g., a wine glass falling down will break when it hits the ground. How these expectations affect relatively low-level perceptual decisions, however, has not been systematically studied previously. To seek an answer to this question, we conducted a behavioral experiment using animations of various familiar objects (e.g., key, wine glass, etc.) freely falling and hitting the ground. During a training session, participants first built expectations about the dynamic properties of those objects. Half of the participants (N = 28) built expectations consistent with their daily lives (e.g., a key bounces rigidly), whereas the other half learned an atypical behavior (e.g., a key wobbles). This was followed by experimental sessions, in which expectations were unmet in 20% of the trials. In both training and experimental sessions, the participant's task was to report whether the objects broke or not upon hitting the ground. Critically, a specific object always remained intact or broke - only the manner in which it did so differed. For example, a key could wobble or remain rigid but never break. We found that participants' reaction times were longer when expectations were unmet, not only for typical material behavior but also when those expectations were atypical and learned during the training session. Furthermore, we found an interplay between long-term and newly learned expectations. Overall, our results show that expectations about material properties can impact relatively low-level perceptual decision-making processes.

Tilt aftereffect spreads across the visual field.

The tilt aftereffect (TAE) is observed when adaptation to a tilted contour alters the perceived tilt of a subsequently presented contour. Thus far, TAE has been treated as a local aftereffect observed only at the location of the adapter. Whether and how TAE spreads to other locations in the visual field has not been systematically studied. Here, we sought an answer to this question by measuring TAE magnitudes at locations including but not limited to the adapter location. The adapter was a tilted grating presented at the same peripheral location throughout an experimental session. In a single trial, participants indicated the perceived tilt of a test grating presented after the adapter at one of fifteen locations in the same visual hemifield as the adapter. We found non-zero TAE magnitudes in all locations tested, showing that the effect spreads across the tested visual hemifield. Next, to establish a link between neuronal activity and behavioral results and to predict the possible neuronal origins of the spread, we built a computational model based on known characteristics of the visual cortex. The simulation results showed that the model could successfully capture the pattern of the behavioral results. Furthermore, the pattern of the optimized receptive field sizes suggests that mid-level visual areas, such as V4, could be critically involved in TAE and its spread across the visual field.

Increasing the spatial extent of attention strengthens surround suppression.

Here we investigate how the extent of spatial attention affects center-surround interaction in visual motion processing. To do so, we measured motion direction discrimination thresholds in humans using drifting gratings and two attention conditions. Participants were instructed to limit their attention to the central part of the stimulus under the narrow attention condition, and to both central and surround parts under the wide attention condition. We found stronger surround suppression under the wide attention condition. The magnitude of the attention effect increased with the size of the surround when the stimulus had low contrast, but did not change when it had high contrast. Results also showed that attention had a weaker effect when the center and surround gratings drifted in opposite directions. Next, to establish a link between the behavioral results and the neuronal response characteristics, we performed computer simulations using the divisive normalization model. Our simulations showed that using smaller versus larger multiplicative attentional gain and parameters derived from the medial temporal (MT) area of the cortex, the model can successfully predict the observed behavioral results. These findings reveal the critical role of spatial attention on surround suppression and establish a link between neuronal activity and behavior. Further, these results also suggest that the reduced surround suppression found in certain clinical disorders (e.g., schizophrenia and autism spectrum disorder) may be caused by abnormal attention mechanisms.

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.

Unmet expectations delay sensory processes.

Expectations strongly affect and shape our perceptual decision-making processes. Specifically, valid expectations speed up perceptual decisions, and determine what we see in a noisy stimulus. Despite the well-established effects of expectations on decision-making, whether and how they affect low-level sensory processes remain elusive. To address this problem, we investigated the effect of expectation on temporal thresholds in an individuation task (detection of the position of an intact image, a house or face). We found that compared to a neutral baseline, thresholds increase when the intact images are of the unexpected category, but remain unchanged when they are of the expected category. Using a recursive Bayesian model with dynamic priors we show that delay in sensory processes is the result of further processing, consequently longer time, required in case of violated expectations. Expectations, however, do not alter internal parameters of the system. These results reveal that sensory processes are delayed when expectations are not met, and a simple parsimonious computational model can successfully explain this effect.

Cortical connectivity in the face of congenital structural changes-A case of homozygous LAMC3 mutation.

The homozygous LAMC3 gene mutation is associated with severe bilateral smoothening and thickening of the lateral occipital cortex . Despite this and further significant changes in gray matter structure, a patient harboring this mutation exhibited a range of remarkably intact perceptual abilities . One possible explanation of this perceptual sparing could be that the white matter structural integrity and functional connectivity in relevant pathways remained intact. To test this idea, we used diffusion tensor and functional magnetic resonance imaging to investigate functional connectivity in resting-state networks in major structural pathways involved in object perception and visual attention and corresponding microstructural integrity in a patient with homozygous LAMC3 mutation and sex, age, education, and socioeconomically matched healthy control group. White matter microstructural integrity results indicated widespread disruptions in both intra- and interhemispheric structural connections except inferior longitudinal fasciculus. With a few exceptions, the functional connectivity between the patient's adjacent gray matter regions of major white matter tracts of interest was conserved. In addition, functional localizers for face, object, and place areas showed similar results with a representative control, providing an explanation for the patient's intact face, place, and object recognition abilities. To generalize this finding, we also compared functional connectivity between early visual areas and face, place, and object category-selective areas, and we found that the functional connectivity of the patient was not different from the control group. Overall, our results provided complementary information about the effects of LAMC3 gene mutation on the human brain including intact temporo-occipital structural and functional connectivity that are compatible with preserved perceptual abilities.

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.

Responses to lightness variations in early human visual cortex.

Lightness is the apparent reflectance of a surface, and it depends not only on the actual luminance of the surface but also on the context in which the surface is viewed [1-10]. The cortical mechanisms of lightness processing are largely unknown, and the role of early cortical areas is still a matter of debate [11-17]. We studied the cortical responses to lightness variations in early stages of the human visual system with functional magnetic resonance imaging (fMRI) while observers were performing a demanding fixation task. The set of dynamically presented visual stimuli included the rectangular version of the classic Craik-O'Brien stimulus [3, 18, 19] and a variant that led to a weaker lightness effect, as well as a pattern with actual luminance variations. We found that the cortical activity in retinotopic areas, including the primary visual cortex (V1), is correlated with context-dependent lightness variations.

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.

Estimating the glossiness transfer function induced by illumination change and testing its transitivity.

The light reflected from a glossy surface depends on the reflectance properties of that surface as well as the flow of light in the scene, the light field. We asked four observers to compare the glossiness of pairs of surfaces under two different real-word light fields, and used this data to estimate a transfer function that captures how perceived glossiness is remapped in changing from one real-world light field to a second. We wished to determine the form of the transfer function and to test whether for any set of three light fields the transfer function from light field 1 to light field 2 and the transfer function from light field 2 to light field 3 could be used to predict the glossiness transfer function from light field 1 to light field 3. Observers' estimated glossiness transfer functions for three sets of light fields were best described by a linear model. The estimated transfer functions exhibited the expected transitivity pattern for three out of four observers. The failure of transitivity for one observer, while significant, was less than 12.5% of the gloss range.

Perceptual grouping-dependent lightness processing in human early visual cortex.

Lightness, the perceived relative achromatic reflectance of a surface, depends strongly on the context within which the surface is viewed. Modest changes in the two-dimensional configuration or three-dimensional scene geometry may lead to profound variations in lightness even though the surface luminance remains constant. Despite recent progress, we are far from a complete understanding of how various aspects of spatial context affect lightness processing in the cortex. Here we use a novel stimulus to show that perceptual grouping through occluders can affect lightness. We first report behavioral results showing how lightness across occlusion depends on spatially distant image features, including luminance and contrast. Next using functional magnetic resonance imaging (fMRI) we show that human early visual cortex responds strongly to occlusion-dependent lightness variations with little or no attention. These results suggest that elements of three-dimensional scene interpretation play a role in early cortical processing of lightness.

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.