Local contour features contribute to figure-ground segregation in monkey V4 neural populations and human perception.

Figure-ground (FG) segregation is a crucial step towards the recognition of objects in natural scenes. Gestalt psychologists have emphasized the importance of contour features in perception of FG. Recent electrophysiological studies have identified a neural population in V4 that shows FG-dependent modulation (FG neurons).

Representation of Natural Contours by a Neural Population in Monkey V4.

The cortical visual area, V4, has been considered to code contours that contribute to the intermediate-level representation of objects. The neural responses to the complex contour features intrinsic to natural contours are expected to clarify the essence of the representation. To approach the cortical coding of natural contours, we investigated the simultaneous coding of multiple contour features in monkey () V4 neurons and their population-level representation.

Correlation between neural responses and human perception in figure-ground segregation.

Segmentation of a natural scene into objects (figures) and background (ground) is one of crucial functions for object recognition and scene understanding. Recent studies have investigated neural mechanisms underlying figure-ground (FG) segregation and reported neural modulation to FG in the intermediate-level visual area, V4, of macaque monkeys (FG neurons). However, whether FG neurons contribute to the perception of FG segregation has not been clarified.

Dimensionality of the intermediate-level representation of shape and texture in monkey V4.

The visual area V4 has been considered to play a crucial role in the intermediate representation of objects, where low-level image features are transformed into object-level representations. We estimated the intrinsic dimensionality in V4 for the representation of local patches generated from natural scenes. The dimensionality was approximately 40, which is approximately half of that reported in IT for the representation of whole natural objects.

Figure-ground responsive fields of monkey V4 neurons estimated from natural image patches.

Neurons in visual area V4 modulate their responses depending on the figure-ground (FG) organization in natural images containing a variety of shapes and textures. To clarify whether the responses depend on the extents of the figure and ground regions in and around the classical receptive fields (CRFs) of the neurons, we estimated the spatial extent of local figure and ground regions that evoked FG-dependent responses (RF-FGs) in natural images and their variants. Specifically, we applied the framework of spike triggered averaging (STA) to the combinations of neural responses and human-marked segmentation images (FG labels) that represent the extents of the figure and ground regions in the corresponding natural image stimuli.

Interaction of surface pattern and contour shape in the tilt after effects evoked by symmetry.

Integration of multiple properties of an object is a fundamental function of the visual cortex in object recognition. For instance, surface patterns and contour shapes are thought to be crucial characteristics that jointly contribute to recognition. However, the mechanisms of integration and corresponding cortical representations have not been fully clarified.

Population coding of figure and ground in natural image patches by V4 neurons.

Segmentation of a natural scene into objects and background is a fundamental but challenging task for recognizing objects. Investigating intermediate-level visual cortical areas with a focus on local information is a crucial step towards understanding the formation of the cortical representations of figure and ground. We examined the activity of a population of macaque V4 neurons during the presentation of natural image patches and their respective variations.

Invariance to low-level features and partial transfer over space in the tilt aftereffects evoked by symmetrical patterns.

The symmetry axis is the midline that divides a pattern into congruent halves, which is physically nonexistent but evokes tilt aftereffect (TAE). To investigate the cortical correspondence of the symmetry axis, we examined the invariance of symmetry-induced TAE with regard to low-level visual features and the spatial transfer of TAE over visual fields. When the adaptation pattern was rotated and changed sequentially with the orientation of the symmetry axis unchanged, the measured TAE decreased only slightly (18%) compared to stationary patterns.

Interactions Elicited by the Contradiction Between Figure Direction Discrimination and Figure-Ground Segregation.

Figure-ground (FG) segregation that separates an object from the rest of the image is a fundamental problem in vision science. A majority of neurons in monkey V2 showed the selectivity to border ownership (BO) that indicates which side of a contour owns the border. Although BO could be a precursor of FG segregation, the contribution of BO to FG segregation has not been clarified.

Modeling the Time-Course of Responses for the Border Ownership Selectivity Based on the Integration of Feedforward Signals and Visual Cortical Interactions.

Border ownership (BO) indicates which side of a contour owns a border, and it plays a fundamental role in figure-ground segregation. The majority of neurons in V2 and V4 areas of monkeys exhibit BO selectivity. A physiological work reported that the responses of BO-selective cells show a rapid transition when a presented square is flipped along its classical receptive field (CRF) so that the opposite BO is presented, whereas the transition is significantly slower when a square with a clear BO is replaced by an ambiguous edge, e.

Sparse coding generates curvature selectivity in V4 neurons.

The cortical area V4 produces a representation of curvature as the intermediate-level representation of an object's shape. We investigated whether sparse coding is the principle driving the generation of the spatial properties of the receptive field in V4 that exhibit curvature selectivity. To investigate the role of sparseness in the construction of curvature representations, we applied component analysis with a sparseness constraint to the activity of model V2 neurons that were responding to shapes derived from natural images.

Perceptual representation and effectiveness of local figure-ground cues in natural contours.

A contour shape strongly influences the perceptual segregation of a figure from the ground. We investigated the contribution of local contour shape to figure-ground segregation. Although previous studies have reported local contour features that evoke figure-ground perception, they were often image features and not necessarily perceptual features.

Facilitatory mechanisms of specular highlights in the perception of depth.

We investigated whether specular highlights facilitate the perception of shape from shading in a search paradigm and how highlights interact with shading to facilitate this perception. Our results indicated that stimuli containing highlights led to shorter searching time with the dependence on the light source direction (top lights make searching faster), suggesting that highlights indeed facilitate shape-from-shading processing. To examine how highlight processing interacts with shading processing, we tested unnatural stimuli for which the lighting directions for shading and highlights were inconsistent.

Early representation of shape by onset synchronization of border-ownership-selective cells in the V1-V2 network.

Construction of surface is a crucial step toward the representation of shape through the integration of local information. Physiological studies have reported that the primary visual cortex (V1) codes the medial axis (MA) that is a skeletal structure equidistant from nearby contours, suggesting the early representation of surface in V1. Although the neural basis of surface construction has not been clarified, the onset synchronization of border ownership (BO)-selective cells is a plausible candidate for the generation of surface.

Spatial and feature-based attention in a layered cortical microcircuit model.

Directing attention to the spatial location or the distinguishing feature of a visual object modulates neuronal responses in the visual cortex and the stimulus discriminability of subjects. However, the spatial and feature-based modes of attention differently influence visual processing by changing the tuning properties of neurons. Intriguingly, neurons' tuning curves are modulated similarly across different visual areas under both these modes of attention.

Feature-based attention in early vision for the modulation of figure-ground segregation.

We investigated psychophysically whether feature-based attention modulates the perception of figure-ground (F-G) segregation and, based on the results, we investigated computationally the neural mechanisms underlying attention modulation. In the psychophysical experiments, the attention of participants was drawn to a specific motion direction and they were then asked to judge the side of figure in an ambiguous figure with surfaces consisting of distinct motion directions. The results of these experiments showed that the surface consisting of the attended direction of motion was more frequently observed as figure, with a degree comparable to that of spatial attention (Wagatsuma et al.

Consistent and robust determination of border ownership based on asymmetric surrounding contrast.

Determination of the figure region in an image is a fundamental step toward surface construction, shape coding, and object representation. Localized, asymmetric surround modulation, reported neurophysiologically in early-to-intermediate-level visual areas, has been proposed as a mechanism for figure-ground segregation. We investigated, computationally, whether such surround modulation is capable of yielding consistent and robust determination of figure side for various stimuli.

Decoding of depth and motion in ambiguous binocular perception.

We have shown previously that random dots with an interocular time delay (ITD), the time difference of the onset of dots between the two eyes, yield both apparent depth and motion, although depth and velocity are covariant and, thus, ITD is inherently ambiguous. The depth of random dots with ITD was proportional to ITD, suggesting that the visual system assumes a constant velocity of the dots and determines depth on the basis of this constant velocity. We performed psychophysical experiments to investigate whether subjects perceive a constant velocity with a variety of ITDs in random dots aligned along a single vertical line that ensures neither apparent motion nor accidental disparity between the dots.

Spatial attention in early vision for the perception of border ownership.

Spatial attention alters contrast gain in early visual areas, which might affect the determination of border ownership (BO) that indicates the direction of figure with respect to the border. We investigated the role of spatial attention applied to early vision in the determination of BO with a computational model that consists of V1, V2, and posterior parietal (PP) modules. Attention alters contrast gain in the V1 module so that it enhances local contrast.

Surround modulation in visual cortex can predict border-ownership selectivity: psychophysical study of border-ownership-dependent tilt aftereffect.

Recent physiological studies revealed that neurons in the macaque visual cortex encode the direction of a figure along a contour (border ownership, BO). Although their cortical mechanisms have not been clarified, a computational model for BO has suggested that surround modulation in early vision can play an important role. Here we examined psychophysically how the strength of BO-dependent tilt aftereffect (BO-TAE) is modulated by a stimulus outside the adapted location in relation to the strength of surround modulation reported in physiological experiments.

Border-ownership-dependent tilt aftereffect in incomplete figures.

A recent physiological finding of neural coding for border ownership (BO) that defines the direction of a figure with respect to the border has provided a possible basis for figure-ground segregation. To explore the underlying neural mechanisms of BO, we investigated stimulus configurations that activate BO circuitry through psychophysical investigation of the BO-dependent tilt aftereffect (BO-TAE). Specifically, we examined robustness of the border ownership signal by determining whether the BO-TAE is observed when gestalt factors are broken.

Facilitation of shape-from-shading perception by random textures.

Most natural objects have a texture on their surface, so the segregation between shading and texture is crucial for the robust perception of three-dimensional structure: The visual system has to decide whether shading or texture evoked the luminance change. We found that the contextual pop-out that results from shading was not suppressed, but was even facilitated, when random texture was added to the luminance of the entire stimulus, indicating the functional segregation and facilitative interaction between shading and texture cues. The local contrast evoked by random texture within a figure or at a boundary was a major factor in the facilitation, suggesting the crucial role of early vision in the interaction between the cues.

Surrounding suppression and facilitation in the determination of border ownership.

Contextual modulation reported in early- to intermediate-level visual areas could be an essential component to signal border ownership (BO) that specifies the direction of figure along a contour. The surrounding regions that evoke significant suppression or facilitation are highly localized and asymmetric with respect to the center of the classical receptive field (CRF). We propose a hypothesis that such surrounding modulation is a basis for BO-selectivity.

Perception of depth and motion from ambiguous binocular information.

The visual system can determine motion and depth from ambiguous information contained in images projected onto both retinas over space and time. The key to the way the system overcomes such ambiguity lies in dependency among multiple cues--such as spatial displacement over time, binocular disparity, and interocular time delay--which might be established based on prior knowledge or experience, and stored in spatiotemporal response characteristics of neurons at an early cortical stage. We conducted a psychophysical investigation of whether a single ambiguous cue (specifically, interocular time delay) permits depth discrimination and motion perception.

Neural grouping and geometric effect in the determination of apparent orientation.

We propose that neural grouping of retinotopically distributed responses in the primary visual cortex (V1) is essential for the determination of apparent tilt, including the tilt illusion. Our psychophysical study shows that apparent tilt is independent of stereo disparity, hue, or contrast of bars, which determine the ownership of their intersection. This leads us to suspect that the neuronal responses within the intersection are excluded from the computation of apparent tilt.