Context modulates the ERP signature of contour integration

We investigated how the electrophysiological signature of contour integration is changed by the context in which a contour is embedded. Specifically, we manipulated the orientations of Gabor elements surrounding an embedded shape outline. The amplitudes of early visual components over posterior scalp regions were changed by the presence of a contour, and by the orientation of elements surrounding the contour. Differences in context type had an effect on the early P1 and N1 components, but not on the later P2 component. The presence of a contour had an effect on the N1 and P2 components, but not on the earlier P1 component. A modulatory effect of context on contour integration was observed on the N1 component. These results highlight the importance of the context in which contour integration takes place.     

Phase-locked alpha oscillations evoked by illusory contour perception in normal and autistic preschool boys

We examined temporal dynamics of EEG phase-locked alpha oscillations during perception of illusory (Kanizsa square) and non-illusory images in boys with autism and age-matched typically developing boys. In typically developing boys the illusory contour (IC) as compared to the control stimulus provoked an increased alpha response at the parietal scalp areas. This IC effect demonstrated continuity within the time window of 133-267 ms after the stimulus onset. Although boys with autism did not display this effect at the group level, part of the sample showed an atypical two-stage pattern of illusory contour effect. The first early stage of IC effect (50-133 ms) was pronounced at the midline occipital electrode localized in the vicinity of the primary visual cortex. The localization and the early onset time suggest that this early IC effect is related to abnormally enhanced "low-level" locally-oriented processes of contour completion in autism. The second stage of IC effect (267-400 ms) was observed at the left parietal region only, and was delayed comparatively to that in healthy boys, suggesting the deficit of "intermediate" processes of perceptual grouping linked to the higher-order visual areas.     

Perceptual organization of local elements into global shapes in the human visual cortex

The question of how local image features on the retina are integrated into perceived global shapes is central to our understanding of human visual perception. Psychophysical investigations have suggested that the emergence of a coherent visual percept, or a "good-Gestalt", is mediated by the perceptual organization of local features based on their similarity. However, the neural mechanisms that mediate unified shape perception in the human brain remain largely unknown. Using human fMRI, we demonstrate that not only higher occipitotemporal but also early retinotopic areas are involved in the perceptual organization and detection of global shapes. Specifically, these areas showed stronger fMRI responses to global contours consisting of collinear elements than to patterns of randomly oriented local elements. More importantly, decreased detection performance and fMRI activations were observed when misalignment of the contour elements disturbed the perceptual coherence of the contours. However, grouping of the misaligned contour elements by disparity resulted in increased performance and fMRI activations, suggesting that similar neural mechanisms may underlie grouping of local elements to global shapes by different visual features (orientation or disparity). Thus, these findings provide novel evidence for the role of both early feature integration processes and higher stages of visual analysis in coherent visual perception.     

Top-Down Control in Contour Grouping

Human observers tend to group oriented line segments into full contours if they follow the Gestalt rule of ''good continuation''. It is commonly assumed that contour grouping emerges automatically in early visual cortex. In contrast, recent work in animal models suggests that contour grouping requires learning and thus involves top-down control from higher brain structures. Here we explore mechanisms of top-down control in perceptual grouping by investigating synchronicity within EEG oscillations. Human participants saw two micro-Gabor arrays in a random order, with the task to indicate whether the first (S1) or the second stimulus (S2) contained a contour of collinearly aligned elements. Contour compared to non-contour S1 produced a larger posterior post-stimulus beta power (15–21 Hz). Contour S2 was associated with a pre-stimulus decrease in posterior alpha power (11–12 Hz) and in fronto-posterior theta (4–5 Hz) phase couplings, but not with a post-stimulus increase in beta power. The results indicate that subjects used prior knowledge from S1 processing for S2 contour grouping. Expanding previous work on theta oscillations, we propose that long-range theta synchrony shapes neural responses to perceptual groupings regulating lateral inhibition in early visual cortex.     

Radial Frequency Analysis of Contour Shapes in the Visual Cortex

Cumulative psychophysical evidence suggests that the shape of closed contours is analysed by means of their radial frequency components (RFC). However, neurophysiological evidence for RFC-based representations is still missing. We investigated the representation of radial frequency in the human visual cortex with functional magnetic resonance imaging. We parametrically varied the radial frequency, amplitude and local curvature of contour shapes. The stimuli evoked clear responses across visual areas in the univariate analysis, but the response magnitude did not depend on radial frequency or local curvature. Searchlight-based, multivariate representational similarity analysis revealed RFC specific response patterns in areas V2d, V3d, V3AB, and IPS0. Interestingly, RFC-specific representations were not found in hV4 or LO, traditionally associated with visual shape analysis. The modulation amplitude of the shapes did not affect the responses in any visual area. Local curvature, SF-spectrum and contrast energy related representations were found across visual areas but without similar specificity for visual area that was found for RFC. The results suggest that the radial frequency of a closed contour is one of the cortical shape analysis dimensions, represented in the early and mid-level visual areas.     

Seeing more than meets the eye: processing of illusory contours in animals

This review article illustrates that mammals, birds and insects are able to perceive illusory contours. Illusory contours lack a physical counterpart, but monkeys, cats, owls and bees perceive them as if they were real borders. In all of these species, a neural correlate for such perceptual completion phenomena has been described. The robustness of neuronal responses and the abundance of cells argue that such neurons might indeed represent a neural correlate for illusory contour perception. The internal state of an animal subject (i.e., alert and behaving) seems to be an important factor when correlating neural activity with perceptual phenomena. The fact that the neural network necessary for illusory contour perception has been found in relatively early visual brain areas in all tested animals suggests that bottom-up processing is largely sufficient to explain such perceptual abilities. However, recent findings in monkeys indicate that feedback loops within the visual system may provide additional modulation. The detection of illusory contours by independently evolved visual systems argues that processing of edges in the absence of contrast gradients reflects fundamental visual constraints and not just an artifact of visual processing.     

Cortical connections and early visual function: intra- and inter-columnar processing.

While it is widely assumed that the long-range horizontal connections in V1 are present to support contour integration, there has been only limited consideration of other possible relationships between anatomy and physiology (the horizontal connections) and visual function beyond contour integration. We introduce the possibility of other relationships directly from the perspective of computation and differential geometry by identifying orientation columns in visual physiology with the (unit) tangent bundle in differential geometry. This suggests abstracting early vision in a space that incorporates both position and orientation, from which we show that the physiology is capable of supporting a number of functional computations beyond contour integration, including texture-flow and shading-flow integration, as well as certain relationships between them. The geometric abstraction emphasizes the role of curvature, which necessitates a coupled investigation into how it might be estimated. The result is an elaboration of layer-to-layer interactions within an orientation column, with non-linearities possibly implemented by shunting inhibition. Finally, we show how the same computational framework naturally lends itself to solving stereo correspondence, with binocular tangents abstracting curves in space.     

A sparse object coding scheme in area V4

Sparse coding has long been recognized as a primary goal of image transformation in the visual system. Sparse coding in early visual cortex is achieved by abstracting local oriented spatial frequencies and by excitatory/inhibitory surround modulation. Object responses are thought to be sparse at subsequent processing stages, but neural mechanisms for higher-level sparsification are not known. Here, convergent results from macaque area V4 neural recording and simulated V4 populations trained on natural object contours suggest that sparse coding is achieved in midlevel visual cortex by emphasizing representation of acute convex and concave curvature. We studied 165 V4 neurons with a random, adaptive stimulus strategy to minimize bias and explore an unlimited range of contour shapes. V4 responses were strongly weighted toward contours containing acute convex or concave curvature. In contrast, the tuning distribution in nonsparse simulated V4 populations was strongly weighted toward low curvature. But as sparseness constraints increased, the simulated tuning distribution shifted progressively toward more acute convex and concave curvature, matching the neural recording results. These findings indicate a sparse object coding scheme in midlevel visual cortex based on uncommon but diagnostic regions of acute contour curvature.     

基于初级视觉皮层的图像匹配模型

提出一种基于初级视觉皮层的图像匹配模型。该模型只采用方位选择性细胞和皮层内有限范围水平连接等Ｖ１基本单元,它以链码表示的目标轮廓作为知识,允许该知识以时间脉冲的形式控制Ｖ１区内神经细胞的动态活动,使与知识轮廓形状相符合的轮廓内的细胞,逐步进入并维持在兴奋状态,最终实现对视野中特定目标轮廓的提取     

Computational modeling and exploration of contour integration for visual saliency

We propose a computational model of contour integration for visual saliency. The model uses biologically plausible devices to simulate how the representations of elements aligned collinearly along a contour in an image are enhanced. Our model adds such devices as a dopamine-like fast plasticity, local GABAergic inhibition and multi-scale processing of images. The fast plasticity addresses the problem of how neurons in visual cortex seem to be able to influence neurons they are not directly connected to, for instance, as observed in contour closure effect. Local GABAergic inhibition is used to control gain in the system without using global mechanisms which may be non-plausible given the limited reach of axonal arbors in visual cortex. The model is then used to explore not only its validity in real and artificial images, but to discover some of the mechanisms involved in processing of complex visual features such as junctions and end-stops as well as contours. We present evidence for the validity of our model in several phases, starting with local enhancement of only a few collinear elements. We then test our model on more complex contour integration images with a large number of Gabor elements. Sections of the model are also extracted and used to discover how the model might relate contour integration neurons to neurons that process end-stops and junctions. Finally, we present results from real world images. Results from the model suggest that it is a good current approximation of contour integration in human vision. As well, it suggests that contour integration mechanisms may be strongly related to mechanisms for detecting end-stops and junction points. Additionally, a contour integration mechanism may be involved in finding features for objects such as faces. This suggests that visual cortex may be more information efficient and that neural regions may have multiple roles.     

Integration of local features into global shapes: monkey and human FMRI studies

The integration of local image features into global shapes was investigated in monkeys and humans using fMRI. An adaptation paradigm was used, in which stimulus selectivity was deduced by changes in the course of adaptation of a pattern of randomly oriented elements. Accordingly, we observed stronger activity when orientation changes in the adapting stimulus resulted in a collinear contour than a different random pattern. This selectivity to collinear contours was observed not only in higher visual areas that are implicated in shape processing, but also in early visual areas where selectivity depended on the receptive field size. These findings suggest that unified shape perception in both monkeys and humans involves multiple visual areas that may integrate local elements to global shapes at different spatial scales.     

Abnormal Contextual Modulation of Visual Contour Detection in Patients with Schizophrenia

Schizophrenia patients demonstrate perceptual deficits consistent with broad dysfunction in visual context processing. These include poor integration of segments forming visual contours, and reduced visual contrast effects (e.g. weaker orientation-dependent surround suppression, ODSS). Background image context can influence contour perception, as stimuli near the contour affect detection accuracy. Because of ODSS, this contextual modulation depends on the relative orientation between the contour and flanking elements, with parallel flankers impairing contour perception. However in schizophrenia, the impact of abnormal ODSS during contour perception is not clear. It is also unknown whether deficient contour perception marks genetic liability for schizophrenia, or is strictly associated with clinical expression of this disorder. We examined contour detection in 25 adults with schizophrenia, 13 unaffected first-degree biological relatives of schizophrenia patients, and 28 healthy controls. Subjects performed a psychophysics experiment designed to quantify the effect of flanker orientation during contour detection. Overall, patients with schizophrenia showed poorer contour detection performance than relatives or controls. Parallel flankers suppressed and orthogonal flankers enhanced contour detection performance for all groups, but parallel suppression was relatively weaker for schizophrenia patients than healthy controls. Relatives of patients showed equivalent performance with controls. Computational modeling suggested that abnormal contextual modulation in schizophrenia may be explained by suppression that is more broadly tuned for orientation. Abnormal flanker suppression in schizophrenia is consistent with weaker ODSS and/or broader orientation tuning. This work provides the first evidence that such perceptual abnormalities may not be associated with a genetic liability for schizophrenia.     

End-stopping and the aperture problem: two-dimensional motion signals in macaque V1

Our perception of fine visual detail relies on small receptive fields at early stages of visual processing. However, small receptive fields tend to confound the orientation and velocity of moving edges, leading to ambiguous or inaccurate motion measurements (the aperture problem). Thus, it is often assumed that neurons in primary visual cortex (V1) carry only ambiguous motion information. Here we show that a subpopulation of V1 neurons is capable of signaling motion direction in a manner that is independent of contour orientation. Specifically, end-stopped V1 neurons obtain accurate motion measurements by responding only to the endpoints of long contours, a strategy which renders them largely immune to the aperture problem. Furthermore, the time course of end-stopping is similar to the time course of motion integration by MT neurons. These results suggest that cortical neurons might represent object motion by responding selectively to two-dimensional discontinuities in the visual scene.     

Ensemble Empirical Mode Decomposition Analysis of EEG Data Collected during a Contour Integration Task

We discuss a data-driven analysis of EEG data recorded during a combined EEG/fMRI study of visual processing during a contour integration task. The analysis is based on an ensemble empirical mode decomposition (EEMD) and discusses characteristic features of event related modes (ERMs) resulting from the decomposition. We identify clear differences in certain ERMs in response to contour vs noncontour Gabor stimuli mainly for response amplitudes peaking around 100 [ms] (called P100) and 200 [ms] (called N200) after stimulus onset, respectively. We observe early P100 and N200 responses at electrodes located in the occipital area of the brain, while late P100 and N200 responses appear at electrodes located in frontal brain areas. Signals at electrodes in central brain areas show bimodal early/late response signatures in certain ERMs. Head topographies clearly localize statistically significant response differences to both stimulus conditions. Our findings provide an independent proof of recent models which suggest that contour integration depends on distributed network activity within the brain.     

Transgenically encoded protein photoinactivation (FlAsH-FALI): acute inactivation of synaptotagmin I

Perceptual completion can link widely separated contour fragments and interpolate illusory contours (ICs) between them. The mechanisms underlying such long-range linking are not well understood. Here we report that completion is much poorer when ICs cross the vertical meridian than when they reside entirely within the left or right visual hemifield. This deficit reflects limitations in cross-hemispheric integration. We also show that the sensitivity to the interhemispheric divide is unique to perceptual completion: a comparable task which did not require completion showed no across-meridian impairment. We propose that these findings support the existence of specialized completion mechanisms in early visual cortical areas (V1/V2), since those areas are likely to be more sensitive to the interhemispheric divide.     

Contour saliency in primary visual cortex

Contour integration is an important intermediate stage of object recognition, in which line segments belonging to an object boundary are perceptually linked and segmented from complex backgrounds. Contextual influences observed in primary visual cortex (V1) suggest the involvement of V1 in contour integration. Here, we provide direct evidence that, in monkeys performing a contour detection task, there was a close correlation between the responses of V1 neurons and the perceptual saliency of contours. Receiver operating characteristic analysis showed that single neuronal responses encode the presence or absence of a contour as reliably as the animal's behavioral responses. We also show that the same visual contours elicited significantly weaker neuronal responses when they were not detected in the detection task, or when they were unattended. Our results demonstrate that contextual interactions in V1 play a pivotal role in contour integration and saliency.     

Learning to link visual contours

In complex visual scenes, linking related contour elements is important for object recognition. This process, thought to be stimulus driven and hard wired, has substrates in primary visual cortex (V1). Here, however, we find contour integration in V1 to depend strongly on perceptual learning and top-down influences that are specific to contour detection. In naive monkeys, the information about contours embedded in complex backgrounds is absent in V1 neuronal responses and is independent of the locus of spatial attention. Training animals to find embedded contours induces strong contour-related responses specific to the trained retinotopic region. These responses are most robust when animals perform the contour detection task but disappear under anesthesia. Our findings suggest that top-down influences dynamically adapt neural circuits according to specific perceptual tasks. This may serve as a general neuronal mechanism of perceptual learning and reflect top-down mediated changes in cortical states.     

基于初级视觉皮层的同步振荡目标检测模型

提出一种基于初级视觉皮层的目标检测模型,该模型只采用方位选择性细胞和皮层内水平连接等Ｖ１基本单元,它以链码表示的目标轮廓作为知识,允许该知识以时间脉冲的形式控制Ｖ１区内神经细胞的动态活动,使与知识轮廓形状相符合的轮廓内的细胞进入同步振荡状态,实现对视野中特定目标轮廓的识别。计算机仿真结果表明,在较高级皮层的“知识”控制之下,初级视觉皮层结构上实现简单的目标检测是可行的。     

Orientation-Cue Invariant Population Responses to Contrast-Modulated and Phase-Reversed Contour Stimuli in Macaque V1 and V2

Visual scenes can be readily decomposed into a variety of oriented components, the processing of which is vital for object segregation and recognition. In primate V1 and V2, most neurons have small spatio-temporal receptive fields responding selectively to oriented luminance contours (first order), while only a subgroup of neurons signal non-luminance defined contours (second order). So how is the orientation of second-order contours represented at the population level in macaque V1 and V2? Here we compared the population responses in macaque V1 and V2 to two types of second-order contour stimuli generated either by modulation of contrast or phase reversal with those to first-order contour stimuli. Using intrinsic signal optical imaging, we found that the orientation of second-order contour stimuli was represented invariantly in the orientation columns of both macaque V1 and V2. A physiologically constrained spatio-temporal energy model of V1 and V2 neuronal populations could reproduce all the recorded population responses. These findings suggest that, at the population level, the primate early visual system processes the orientation of second-order contours initially through a linear spatio-temporal filter mechanism. Our results of population responses to different second-order contour stimuli support the idea that the orientation maps in primate V1 and V2 can be described as a spatial-temporal energy map.