Population coding for figure-ground texture segregation in macaque V1 and V4.

Object recognition often involves the brain segregating objects from their surroundings. Neurophysiological studies of figure-ground texture segregation have yielded inconsistent results, particularly on whether V1 neurons can perform figure-ground texture segregation or just detect texture borders. To address this issue from a population perspective, we utilized two-photon calcium imaging to simultaneously record the responses of large samples of V1 and V4 neurons to figure-ground texture stimuli in awake, fixating macaques. The average response changes indicate that V1 neurons mainly detect texture borders, while V4 neurons are involved in figure-ground segregation. However, population analysis (SVM decoding of PCA-transformed neuronal responses) reveal that V1 neurons not only detect figure-ground borders, but also contribute to figure-ground texture segregation, although requiring substantially more principal components than V4 neurons to reach a 75 % decoding accuracy. Individually, V1/V4 neurons showing larger (negative/positive) figure-ground response differences contribute more to figure-ground segregation. But for V1 neurons, the contribution becomes significant only when many principal components are considered. We conclude that V1 neurons participate in figure-ground segregation primarily by defining the figure borders, and the poorly structured figure-ground information V1 neurons carry could be further utilized by V4 neurons to accomplish figure-ground segregation.

Backward masking implicates cortico-cortical recurrent processes in convex figure context effects and cortico-thalamic recurrent processes in resolving figure-ground ambiguity.

Introduction: Previous experiments purportedly showed that image-based factors like convexity were sufficient for figure assignment. Recently, however, we found that the probability of perceiving a figure on the convex side of a central border was only slightly higher than chance for two-region displays and increased with the number of display regions; this increase was observed only when the concave regions were homogeneously colored. These convex figure context effects (CEs) revealed that figure assignment in these classic displays entails more than a response to local convexity. A Bayesian observer replicated the convex figure CEs using both a convexity object prior and a new, homogeneous background prior and made the novel prediction that the classic displays in which both the convex and concave regions were homogeneous were ambiguous during perceptual organization. Methods: Here, we report three experiments investigating the proposed ambiguity and examining how the convex figure CEs unfold over time with an emphasis on whether they entail recurrent processing. Displays were shown for 100 ms followed by pattern masks after ISIs of 0, 50, or 100 ms. The masking conditions were designed to add noise to recurrent processing and therefore to delay the outcome of processes in which they play a role. In Exp. 1, participants viewed two- and eight-region displays with homogeneous convex regions (homo-convex displays; the putatively ambiguous displays). In Exp. 2, participants viewed putatively unambiguous hetero-convex displays. In Exp. 3, displays and masks were presented to different eyes, thereby delaying mask interference in the thalamus for up to 100 ms. Results and discussion: The results of Exps. 1 and 2 are consistent with the interpretation that recurrent processing is involved in generating the convex figure CEs and resolving the ambiguity of homo-convex displays. The results of Exp. 3 suggested that corticofugal recurrent processing is involved in resolving the ambiguity of homo-convex displays and that cortico-cortical recurrent processes play a role in generating convex figure CEs and these two types of recurrent processes operate in parallel. Our results add to evidence that perceptual organization evolves dynamically and reveal that stimuli that seem unambiguous can be ambiguous during perceptual organization.

Temporal correspondence in perceptual organization: Reciprocal interactions between temporal sensitivity and figure-ground segregation.

How do visual representations account for time? Is it the case that they represent time by themselves possessing temporal properties (temporal mirroring) or by atemporal markers/tags (temporal tagging)? This question has been asked for the past 5 decades and more, in neuroscience, philosophy, and psychology. To address this debate, we designed a study to test temporal correspondence. We tested whether a temporal property (flicker frequency) could influence figure-ground segregation, and in turn, reciprocally, whether a figure-ground segregation would alter a temporal property (here, temporal resolution). We manipulated flicker frequency of dots on either side of an ambiguous edge in Experiment 1 and asked participants to indicate the figural region. In Experiment 2, we measured temporal sensitivity using a temporal order judgment (TOJ) task in both figural and ground regions. We showed temporal correspondence by showing specifically that figure-ground segregation depends on flicker frequency differences between two regions in ambiguous displays, where slow-flickering regions are seen as figural (Experiment 1). Reciprocally, in Experiment 2, we showed that participants performed a temporal-order judgment task better when the task had to be performed on a region seen as background compared with the same region seen as a figure. We show how relatively slower flickering regions are seen as figural, and correspondingly, seeing a region as figural is associated with a poorer temporal resolution. Our results collectively allow us to demonstrate a tight temporal correspondence in figure-ground perception, which could be explained using the parvocellular and magnocellular pathways, the two major retino-geniculo-cortical pathways.

BOLD activation on the groundside of figures: More suppression of grounds that competed more for figural status.

A fundamental aspect of object detection is assigning a border to one (figure) side but not the other (ground) side. Figures are shaped; grounds appear shapeless near the figure border. Accumulating evidence supports the view that the mechanism of figure assignment is inhibitory competition with the figure perceived on the winning side. Suppression has been observed on the groundside of figure borders. One prediction is that more suppression will be observed when the groundside competes more for figural status. We tested this prediction by assessing BOLD activation on the groundside of two types of stimuli with articulated borders: AEnov and AEfam stimuli. In both stimulus types, multiple image-based priors (symmetry, closure, small area, enclosure by a larger region) favored the inside as the figure. In AEfam but not AEnov stimuli, the figural prior of familiar configuration present on the outside competes for figural status. Observers perceived the insides of both types of stimuli as novel figures and the outsides as shapeless grounds. Previously, we observed lower BOLD activation in early visual areas representing the grounds of AEfam than AEnov stimuli, although unexpectedly, activation was above baseline. With articulated borders, it can be difficult to exclude figure activation from ground ROIs. Here, our ground ROIs better excluded figure activation; we also added straight-edge (SE) control stimuli and increased the sample size. In early visual areas representing the grounds, we observed lower BOLD activation on the groundside of AEfam than AEnov stimuli and below-baseline BOLD activation on the groundside of SE and AEfam stimuli. These results, indicating that greater suppression is applied to groundsides that competed more for figural status but lost the competition, support a Bayesian model of figure assignment in which proto-objects activated at both low and high levels where image features and familiar configurations are represented, respectively, compete for figural status.

Task set and instructions influence the weight of figural priors: A psychophysical study with extremal edges and familiar configuration.

In figure-ground organization, the figure is defined as a region that is both "shaped" and "nearer." Here we test whether changes in task set and instructions can alter the outcome of the cross-border competition between figural priors that underlies figure assignment. Extremal edge (EE), a relative distance prior, has been established as a strong figural prior when the task is to report "which side is nearer?" In three experiments using bipartite stimuli, EEs competed and cooperated with familiar configuration, a shape prior for figure assignment in a "which side is shaped?" task." Experiment 1 showed small but significant effects of familiar configuration for displays sketching upright familiar objects, although "shaped-side" responses were predominantly determined by EEs. In Experiment 2, instructions regarding the possibility of perceiving familiar shapes were added. Now, although EE remained the dominant prior, the figure was perceived on the familiar-configuration side of the border on a significantly larger percentage of trials across all display types. In Experiment 3, both task set (nearer/shaped) and the presence versus absence of instructions emphasizing that familiar objects might be present were manipulated within subjects. With familiarity thus "primed," effects of task set emerged when EE and familiar configuration favored opposite sides as figure. Thus, changing instructions can modulate the weighing of figural priors for shape versus distance in figure assignment in a manner that interacts with task set. Moreover, we show that the influence of familiar parts emerges in participants without medial temporal lobe/ perirhinal cortex brain damage when instructions emphasize that familiar objects might be present.

Learned value and predictiveness affect gaze but not figure assignment.

Many factors affect figure-ground segregation, but the contributions of attention and reward history to this process is uncertain. We conducted two experiments to investigate whether reward learning influences figure assignment and whether this relationship was mediated by attention. Participants learned to associate certain shapes with a reward contingency: During a learning phase, they chose between two shapes on each trial, with subsets of shapes associated with high-probability win, low-probability win, high-probability loss, and low-probability loss. In a test phase, participants were given a figure-ground task, in which they indicated which of two regions that shared a contour they perceived as the figure (high-probability win and low-probability win shapes were pitted against each other, as were high-probability loss and low-probability loss shapes). The results revealed that participants had learned the reward contingencies and that, following learning, attention was reliably drawn to the optimal stimulus. Despite this, neither reward history nor the resulting attentional allocation influenced figure-ground organization.

The effect of luminance values of the edge between regions on figure-ground perception.

The present study examined whether luminance values of an edge between regions affect figure-ground perception. A dark-gray or light-gray region was presented on the left (or upper) or right (or lower) side of an edge. Observers reported which region was perceived as a figure. The results in the 0.2° edge width condition showed that dark-gray regions with low luminance edges were perceived as a figure significantly more often than dark-gray regions with high luminance edges. The edge luminance effect occurred, partly occurred, and did not occur when the edge widths were 0.2, 0.5, and 0.8° of visual angle, respectively. These results showed that luminance values of the edge between the regions affected figure-ground perception, and that regions with luminance values closer to that of the edge between the regions were perceived as a figure more often, but only when the edge width was narrow. The edge luminance effect occurred less often as the edge width was increased and would be  attributed to similarity grouping between edge and region.

The role of shape recognition in figure/ground perception in infancy.

In this study we sought to determine whether infants, like adults, utilize previous experience to guide figure/ground processing. After familiarization to a shape, 5-month-olds preferentially attended to the side of an ambiguous figure/ground test stimulus corresponding to that shape, suggesting that they were viewing that portion as the figure. Infants' failure to exhibit this preference in a control condition in which both sides of the test stimulus were displayed as figures indicated that the results in the experimental condition were not due to a preference between two figure shapes. These findings demonstrate for the first time that figure/ground processing in infancy is sensitive to top-down influence. Thus, a critical aspect of figure/ground processing is functional early in life.

Correlations Between Hysteretic Categorical and Continuous Judgments of Perceptual Stimuli Supporting a Unified Dynamical Systems Approach to Perception.

We report from two variants of a figure-ground experiment that is known in the literature to involve a bistable perceptual domain. The first variant was conducted as a two-alternative forced-choice experiment and in doing so tested participants on a categorical measurement scale. The second variant involved a Likert scale measure that was considered to represent a continuous measurement scale. The two variants were conducted as a single within-subjects experiment. Measures of bistability operationalized in terms of hysteresis size scores showed significant positive correlations across the two response conditions. The experimental findings are consistent with a dualistic interpretation of self-organizing perceptual systems when they are described on a macrolevel by means of so-called amplitude equations. This is explicitly demonstrated for a Lotka-Volterra-Haken amplitude equation model of task-related brain activity. As a by-product, the proposed dynamical systems perspective also sheds new light on the anchoring problem of producing numerical, continuous judgments.

Temporal dynamics of different cases of bi-stable figure-ground perception.

Segmentation of a visual scene in "figure" and "ground" is essential for perception of the three-dimensional layout of a scene. In cases of bi-stable perception, two distinct figure-ground interpretations alternate over time. We were interested in the temporal dynamics of these alternations, in particular when the same image is presented repeatedly, with short blank periods in-between. Surprisingly, we found that the intermittent presentation of Rubin's classical "face-or-vase" figure, which is frequently taken as a standard case of bi-stable figure-ground perception, often evoked perceptual switches during the short presentations and stabilization was not prominent. Interestingly, bi-stable perception of Kanizsa's anomalous transparency figure did strongly stabilize across blanks. We also found stabilization for the Necker cube, which we used for comparison. The degree of stabilization (and the lack of it) varied across stimuli and across individuals. Our results indicate, against common expectation, that the stabilization phenomenon cannot be generally evoked by intermittent presentation. We argue that top-down feedback factors such as familiarity, semantics, expectation, and perceptual bias contribute to the complex processes underlying the temporal dynamics of bi-stable figure-ground perception.

Spatially rearranged object parts can facilitate perception of intact whole objects.

The familiarity of an object depends on the spatial arrangement of its parts; when the parts are spatially rearranged, they form a novel, unrecognizable configuration. Yet the same collection of parts comprises both the familiar and novel configuration. Is it possible that the collection of familiar parts activates a representation of the intact familiar configuration even when they are spatially rearranged? We presented novel configurations as primes before test displays that assayed effects on figure-ground perception from memories of intact familiar objects. In our test displays, two equal-area regions shared a central border; one region depicted a portion of a familiar object. Previous research with such displays has shown that participants are more likely to perceive the region depicting a familiar object as the figure and the abutting region as its ground when the familiar object is depicted in its upright orientation rather than upside down. The novel primes comprised either the same or a different collection of parts as the familiar object in the test display (part-rearranged and control primes, respectively). We found that participants were more likely to perceive the familiar region as figure in upright vs. inverted displays following part-rearranged primes but not control primes. Thus, priming with a novel configuration comprising the same familiar parts as the upcoming figure-ground display facilitated orientation-dependent effects of object memories on figure assignment. Similar results were obtained when the spatially rearranged collection of parts was suggested on the groundside of the prime's border, suggesting that familiar parts in novel configurations access the representation of their corresponding intact whole object before figure assignment. These data demonstrate that familiar parts access memories of familiar objects even when they are arranged in a novel configuration.

The homogeneity effect on figure/ground perception in infancy.

We examined whether the homogeneity of the two profiles of Rubin's goblet affects figure/ground perception in infants. We modified the two profiles of Rubin's goblet in order to compare figure/ground perception under four test patterns: (1) two profiles painted with horizontal lines (horizontal-line condition), (2) two profiles painted middle gray (uni-color condition), (3) one profile painted light gray and the other dark gray (two-color condition), and (4) a goblet painted with concentric circles (concentric-circles condition). In the horizontal-line condition the homogeneity of the profile was strengthened, and in the two-color condition the homogeneity of the profile was weakened compared to the uni-color condition, which was an original Rubin's goblet. In the concentric-circles condition the homogeneity of the reversed areas of the horizontal-line were strengthened. After infants were familiarized with each Rubin's goblet, the infants were tested on their discrimination between the two profiles and the goblet in the post-familiarization test. In horizontal-line condition, uni-color condition and concentric-circles condition infants showed a novelty preference for the two profiles in the post-familiarization test. On the other hand, in the two-color condition no preference was observed in the post-familiarization test. This means that infants perceived the goblet as figure and the two profiles as ground in the horizontal-line condition, the uni-color condition and the concentric-circles condition. We found that infants could not perceive the goblet area as figure when the homogeneity of the two profiles was weakened. It can be said that figure/ground perception in infancy is not affected by strengthened homogeneity, but is affected by weakened homogeneity.

Out of Mind, Out of Sight: Unexpected Scene Elements Frequently Go Unnoticed Until Primed.

The human visual system employs a sophisticated set of strategies for scanning the environment and directing attention to stimuli that can be expected given the context and a person's past experience. Although these strategies enable us to navigate a very complex physical and social environment, they can also cause highly salient, but unexpected stimuli to go completely unnoticed. To examine the generality of this phenomenon, we conducted eight studies that included 15 different experimental conditions and 1,577 participants in all. These studies revealed that a large majority of participants do not report having seen a woman in the center of an urban scene who was photographed in midair as she was committing suicide. Despite seeing the scene repeatedly, 46 % of all participants failed to report seeing a central figure and only 4.8 % reported seeing a falling person. Frequency of noticing the suicidal woman was highest for participants who read a narrative priming story that increased the extent to which she was schematically congruent with the scene. In contrast to this robust effect of inattentional blindness, a majority of participants reported seeing other peripheral objects in the visual scene that were equally difficult to detect, yet more consistent with the scene. Follow-up qualitative analyses revealed that participants reported seeing many elements that were not actually present, but which could have been expected given the overall context of the scene. Together, these findings demonstrate the robustness of inattentional blindness and highlight the specificity with which different visual primes may increase noticing behavior.

Organização espacial na percepção visual de luminosidade / Spatial organization in visual perception of lightness

Este estudo analisou a influência de variações físicas dos estímulos, com base na organização espacial de figura-fundo criada pela associação dos efeitos ilusórios de contraste simultâneo de luminosidade e de contornos subjetivos. A cada participante, no total de 64, foram apresentadas 160 matrizes de escolha, cada uma composta de um estímulo modelo e quatro estímulos de comparação, devendo ser identificado qual dos quatro estímulos de comparação correspondia ao estímulo modelo. A diferença significativa entre as médias de ajuste visual verificadas para a condição de contorno subjetivo médio e para a condição controle (sem contorno) mostrou que a formação clássica de contornos subjetivos de Kanizsa, quando associada ao efeito de contraste simultâneo de luminosidade, influenciou a percepção de luminosidade dos participantes.(AU)

This study investigated the influence of physical variations of stimuli, based on the spatial organization of figure-ground, which is created by the association of illusory effects of simultaneous lightness contrast and subjective contours. One hundred and sixty matrixes were presented to each participant, among a total of 64. Each matrix was composed of one sample stimulus and four comparison stimuli, and the task was to identify which of the four comparison stimuli corresponded to the sample stimulus. The significant difference between the averages of visual adjustment that were verified for the medium subjective contour and for the control condition (without contour) showed that the classical formation of Kanizsa subjective contours influenced visual perception when associated with the effect of simultaneous lightness contrast.(AU)

Organização espacial na percepção visual de luminosidade / Spatial organization in visual perception of lightness

Este estudo analisou a influência de variações físicas dos estímulos, com base na organização espacial de figura-fundo criada pela associação dos efeitos ilusórios de contraste simultâneo de luminosidade e de contornos subjetivos. A cada participante, no total de 64, foram apresentadas 160 matrizes de escolha, cada uma composta de um estímulo modelo e quatro estímulos de comparação, devendo ser identificado qual dos quatro estímulos de comparação correspondia ao estímulo modelo. A diferença significativa entre as médias de ajuste visual verificadas para a condição de contorno subjetivo médio e para a condição controle (sem contorno) mostrou que a formação clássica de contornos subjetivos de Kanizsa, quando associada ao efeito de contraste simultâneo de luminosidade, influenciou a percepção de luminosidade dos participantes.

This study investigated the influence of physical variations of stimuli, based on the spatial organization of figure-ground, which is created by the association of illusory effects of simultaneous lightness contrast and subjective contours. One hundred and sixty matrixes were presented to each participant, among a total of 64. Each matrix was composed of one sample stimulus and four comparison stimuli, and the task was to identify which of the four comparison stimuli corresponded to the sample stimulus. The significant difference between the averages of visual adjustment that were verified for the medium subjective contour and for the control condition (without contour) showed that the classical formation of Kanizsa subjective contours influenced visual perception when associated with the effect of simultaneous lightness contrast.