Brightness Perception, Illusory Contours and Corticogeniculate Feedback

A neural network model of early visual processing offers an explanation of brightness effects often associated with illusory contours. Top-down feedback from the model's analog of visual cortical complex cells to model lateral geniculate nucleus (LGN) cells are used to enhance contrast at line ends and other areas of boundary discontinuity. The result is an increase in perceived brightness outside a dark line end, akin to what Kennedy (1979) termed "brightness buttons" in his analysis of visual illusions. When several lines form a suitable configuration, as in an Ehrenstein pattern, the perceptual effect of enhanced brightness can be quite strong. Model simulations show the generation of brightness buttons. With the LGN model circuitry embedded in a larger model of preattentive vision, simulations using complex inputs show the interaction of the brightness buttons with real and illusory contours.

Fast Synchronization of Perpetual Grouping in Laminar Visual Cortical Circuits

Perceptual grouping is well-known to be a fundamental process during visual perception, notably grouping across scenic regions that do not receive contrastive visual inputs. Illusory contours are a classical example of such groupings. Recent psychophysical and neurophysiological evidence have shown that the grouping process can facilitate rapid synchronization of the cells that are bound together by a grouping, even when the grouping must be completed across regions that receive no contrastive inputs. Synchronous grouping can hereby bind together different object parts that may have become desynchronized due to a variety of factors, and can enhance the efficiency of cortical transmission. Neural models of perceptual grouping have clarified how such fast synchronization may occur by using bipole grouping cells, whose predicted properties have been supported by psychophysical, anatomical, and neurophysiological experiments. These models have not, however, incorporated some of the realistic constraints on which groupings in the brain are conditioned, notably the measured spatial extent of long-range interactions in layer 2/3 of a grouping network, and realistic synaptic and axonal signaling delays within and across cells in different cortical layers. This work addresses the question: Can long-range interactions that obey the bipole constraint achieve fast synchronization under realistic anatomical and neurophysiological constraints that initially desynchronize grouping signals? Can the cells that synchronize retain their analog sensitivity to changing input amplitudes? Can the grouping process complete and synchronize illusory contours across gaps in bottom-up inputs? Our simulations show that the answer to these questions is Yes.

Spiking Dynamics during Perceptual Grouping in the Laminar Circuits of Visual Cortex

Grouping of collinear boundary contours is a fundamental process during visual perception. Illusory contour completion vividly illustrates how stable perceptual boundaries interpolate between pairs of contour inducers, but do not extrapolate from a single inducer. Neural models have simulated how perceptual grouping occurs in laminar visual cortical circuits. These models predicted the existence of grouping cells that obey a bipole property whereby grouping can occur inwardly between pairs or greater numbers of similarly oriented and co-axial inducers, but not outwardly from individual inducers. These models have not, however, incorporated spiking dynamics. Perceptual grouping is a challenge for spiking cells because its properties of collinear facilitation and analog sensitivity to inducer configurations occur despite irregularities in spike timing across all the interacting cells. Other models have demonstrated spiking dynamics in laminar neocortical circuits, but not how perceptual grouping occurs. The current model begins to unify these two modeling streams by implementing a laminar cortical network of spiking cells whose intracellular temporal dynamics interact with recurrent intercellular spiking interactions to quantitatively simulate data from neurophysiological experiments about perceptual grouping, the structure of non-classical visual receptive fields, and gamma oscillations.

The Role of Edges and Line-Ends in Illusory Contour Formation

Illusory contours can be induced along directions approximately collinear to edges or approximately perpendicular to the ends of lines. Using a rating scale procedure we explored the relation between the two types of inducers by systematically varying the thickness of inducing elements to result; in varying amounts of "edge-like" or "line-like" induction. Inducers for om illusory figures consisted of concentric rings with arcs missing. Observers judged the clarity and brightness of illusory figures as the number of arcs, their thicknesses, and spacings were parametrically varied. Degree of clarity and amount of induced brightness were both found to be inverted-U functions of the number of arcs. These results mandate that any valid model of illusory contour formation must account for interference effects between parallel lines or between those neural units responsible for completion of boundary signals in directions perpendicular to the ends of thin lines. Line width was found to have an effect on both clarity and brightness, a finding inconsistent with those models which employ only completion perpendicular to inducer orientation.

Cortical Dynamics of Feature Binding and Reset: Control of Visual Persistence

An analysis of the reset of visual cortical circuits responsible for the binding or segmentation of visual features into coherent visual forms yields a model that explains properties of visual persistence. The reset mechanisms prevent massive smearing or visual percepts in response to rapidly moving images. The model simulates relationships among psychophysical data showing inverse relations of persistence to flash luminance and duration, greaterr persistence of illusory contours than real contours, a U-shaped temporal function for persistence of illusory contours, a reduction of persistence: due to adaptation with a stimulus of like orientation, an increase or persistence due to adaptation with a stimulus of perpendicular orientation, and an increase of persistence with spatial separation of a masking stimulus. The model suggests that a combination of habituative, opponent, and endstopping mechanisms prevent smearing and limit persistence. Earlier work with the model has analyzed data about boundary formation, texture segregation, shape-from-shading, and figure-ground separation. Thus, several types of data support each model mechanism and new predictions are made.

The Role of Edges and Line-Ends in Illusory Contour Formation

Illusory contours can be induced along direction approximately collinear to edges or approximately perpendicular to the ends of lines. Using a rating scale procedure we explored the relation between the two types of inducers by systematically varying the thickness of inducing elements to result in varying amounts of "edge-like" or "line-like" induction. Inducers for our illusory figures consisted of concentric rings with arcs missing. Observers judged the clarity and brightness of illusory figures as the number of arcs, their thicknesses, and spacings were parametrically varied. Degree of clarity and amount of induced brightness were both found to be inverted-U functions of the number of arcs. These results mandate that any valid model of illusory contour formation must account for interference effects between parallel lines or between those neural units responsible for completion of boundary signals in directions perpendicular to the ends of thin lines. Line width was found to have an efFect on both clarity and brightness, a finding inconsistent with those models which employ only completion perpendicular to inducer orientation.

Boundary, Brightness, and Depth Interactions During Preattentive Representation and Attentive Recognition of Figure and Ground

This article applies a recent theory of 3-D biological vision, called FACADE Theory, to explain several percepts which Kanizsa pioneered. These include 3-D pop-out of an occluding form in front of an occluded form, leading to completion and recognition of the occluded form; 3-D transparent and opaque percepts of Kanizsa squares, with and without Varin wedges; and interactions between percepts of illusory contours, brightness, and depth in response to 2-D Kanizsa images. These explanations clarify how a partially occluded object representation can be completed for purposes of object recognition, without the completed part of the representation necessarily being seen. The theory traces these percepts to neural mechanisms that compensate for measurement uncertainty and complementarity at individual cortical processing stages by using parallel and hierarchical interactions among several cortical processing stages. These interactions are modelled by a Boundary Contour System (BCS) that generates emergent boundary segmentations and a complementary Feature Contour System (FCS) that fills-in surface representations of brightness, color, and depth. The BCS and FCS interact reciprocally with an Object Recognition System (ORS) that binds BCS boundary and FCS surface representations into attentive object representations. The BCS models the parvocellular LGN→Interblob→Interstripe→V4 cortical processing stream, the FCS models the parvocellular LGN→Blob→Thin Stripe→V4 cortical processing stream, and the ORS models inferotemporal cortex.

A Neural Model of 3D Shape-From-Texture: Multiple-Scale Filtering, Boundary Grouping, and Surface Filling-In

A neural model is presented of how cortical areas V1, V2, and V4 interact to convert a textured 2D image into a representation of curved 3D shape. Two basic problems are solved to achieve this: (1) Patterns of spatially discrete 2D texture elements are transformed into a spatially smooth surface representation of 3D shape. (2) Changes in the statistical properties of texture elements across space induce the perceived 3D shape of this surface representation. This is achieved in the model through multiple-scale filtering of a 2D image, followed by a cooperative-competitive grouping network that coherently binds texture elements into boundary webs at the appropriate depths using a scale-to-depth map and a subsequent depth competition stage. These boundary webs then gate filling-in of surface lightness signals in order to form a smooth 3D surface percept. The model quantitatively simulates challenging psychophysical data about perception of prolate ellipsoids (Todd and Akerstrom, 1987, J. Exp. Psych., 13, 242). In particular, the model represents a high degree of 3D curvature for a certain class of images, all of whose texture elements have the same degree of optical compression, in accordance with percepts of human observers. Simulations of 3D percepts of an elliptical cylinder, a slanted plane, and a photo of a golf ball are also presented.

Laminar Cortical Dynamics of Visual Form and Motion Interactions During Coherent Object Motion Perception

How do visual form and motion processes cooperate to compute object motion when each process separately is insufficient? Consider, for example, a deer moving behind a bush. Here the partially occluded fragments of motion signals available to an observer must be coherently grouped into the motion of a single object. A 3D FORMOTION model comprises five important functional interactions involving the brain’s form and motion systems that address such situations. Because the model’s stages are analogous to areas of the primate visual system, we refer to the stages by corresponding anatomical names. In one of these functional interactions, 3D boundary representations, in which figures are separated from their backgrounds, are formed in cortical area V2. These depth-selective V2 boundaries select motion signals at the appropriate depths in MT via V2-to-MT signals. In another, motion signals in MT disambiguate locally incomplete or ambiguous boundary signals in V2 via MT-to-V1-to-V2 feedback. The third functional property concerns resolution of the aperture problem along straight moving contours by propagating the influence of unambiguous motion signals generated at contour terminators or corners. Here, sparse “feature tracking signals” from, e.g., line ends, are amplified to overwhelm numerically superior ambiguous motion signals along line segment interiors. In the fourth, a spatially anisotropic motion grouping process takes place across perceptual space via MT-MST feedback to integrate veridical feature-tracking and ambiguous motion signals to determine a global object motion percept. The fifth property uses the MT-MST feedback loop to convey an attentional priming signal from higher brain areas back to V1 and V2. The model's use of mechanisms such as divisive normalization, endstopping, cross-orientation inhibition, and longrange cooperation is described. Simulated data include: the degree of motion coherence of rotating shapes observed through apertures, the coherent vs. element motion percepts separated in depth during the chopsticks illusion, and the rigid vs. non-rigid appearance of rotating ellipses.

The Perception of Globally Coherent Motion

How do human observers perceive a coherent pattern of motion from a disparate set of local motion measures? Our research has examined how ambiguous motion signals along straight contours are spatially integrated to obtain a globally coherent perception of motion. Observers viewed displays containing a large number of apertures, with each aperture containing one or more contours whose orientations and velocities could be independently specified. The total pattern of the contour trajectories across the individual apertures was manipulated to produce globally coherent motions, such as rotations, expansions, or translations. For displays containing only straight contours extending to the circumferences of the apertures, observers' reports of global motion direction were biased whenever the sampling of contour orientations was asymmetric relative to the direction of motion. Performance was improved by the presence of identifiable features, such as line ends or crossings, whose trajectories could be tracked over time. The reports of our observers were consistent with a pooling process involving a vector average of measures of the component of velocity normal to contour orientation, rather than with the predictions of the intersection-of-constraints analysis in velocity space.