Dissociation between Attention-Dependent and Spatially Specific Illusory Shape Responses within the Topographic Areas of the Posterior Parietal Cortex.

The human visual system consists of multiple topographic maps that extend from the early visual cortex (EVC) along the dorsal and ventral processing streams. Responses to illusory shapes within these maps have been demonstrated in the ventral stream areas, in particular the lateral occipital complex (LOC). Recently, the intraparietal sulcus (IPS) of the dorsal stream has been linked to the processing of illusory shapes defined by motion. It remains unclear whether the topographically organized parietal areas also respond to stationary illusory shapes, which would suggest their generic role in representing illusory content. In the current study we measured brain responses using fMRI while 30 human participants (12 male) observed flickering inducers around the fixation task. The inducers either formed an illusory diamond in the center, a triangle in the left or right hemifield, or were inverted such that no illusory figure was formed. We compared responses of parietal regions IPS0-IPS5 and SPL1 to each illusory figure with the nonillusory condition. To determine the role of attentional modulation on illusory shape responses we manipulated the difficulty of the fixation task. Our results show that all IPS areas responded to illusory shapes. The more posterior areas IPS0-IPS3 additionally displayed a preference toward contralateral shapes, while the more anterior areas IPS4 and IPS5 showed response attenuation with increased task difficulty. We suggest that the IPS can represent illusory content generated not only by moving, but also by stationary stimuli, and that there is a functional dissociation between attention-dependent anterior and spatially specific posterior topographic maps.SIGNIFICANCE STATEMENT The traditional view of the ventral visual pathway being solely responsible for representation of objects has recently been challenged by demonstrating illusory shape representation within the dorsal visual pathway with moving bistable stimuli. Our results provide evidence for the dorsal stream contribution to representing not only moving, but also stationary illusory shapes. Our results also show a functional subdivision along the topographic maps, with spatially specific shape responses in the more posterior, and attention-dependent responses in the more anterior areas. These findings have implications for our understanding of the relationship between attention and grouping in healthy individuals and neuropsychological patients. Furthermore, IPS areas should be considered in theoretical accounts and models of how subjective content is generated in the brain.

Top-Down Feedback Controls the Cortical Representation of Illusory Contours in Mouse Primary Visual Cortex.

Visual systems have evolved to recognize and extract features from complex scenes using limited sensory information. Contour perception is essential to this process and can occur despite breaks in the continuity of neighboring features. Such robustness of the animal visual system to degraded or occluded shapes may also give rise to an interesting phenomenon of optical illusions. These illusions provide a great opportunity to decipher neural computations underlying contour integration and object detection. Kanizsa illusory contours have been shown to evoke responses in the early visual cortex despite the lack of direct receptive field activation. Recurrent processing between visual areas has been proposed to be involved in this process. However, it is unclear whether higher visual areas directly contribute to the generation of illusory responses in the early visual cortex. Using behavior, in vivo electrophysiology, and optogenetics, we first show that the primary visual cortex (V1) of male mice responds to Kanizsa illusory contours. Responses to Kanizsa illusions emerge later than the responses to the contrast-defined real contours in V1. Second, we demonstrate that illusory responses are orientation-selective. Finally, we show that top-down feedback controls the neural correlates of illusory contour perception in V1. Our results suggest that higher-order visual areas may fill in the missing information in the early visual cortex necessary for illusory contour perception.SIGNIFICANCE STATEMENT Perception of the Kanizsa illusory contours is impaired in neurodevelopmental disorders such as schizophrenia, autism, and Williams syndrome. However, the mechanism of the illusory contour perception is poorly understood. Here we describe the behavioral and neural correlates of Kanizsa illusory contours perception in mice, a genetically tractable model system. We show that top-down feedback controls the neural responses to Kanizsa illusion in V1. To our knowledge, this is the first description of the neural correlates of the Kanizsa illusion in mice and the first causal demonstration of their regulation by top-down feedback.

Brain network mechanisms of visual shape completion.

Visual shape completion recovers object shape, size, and number from spatially segregated edges. Despite being extensively investigated, the process's underlying brain regions, networks, and functional connections are still not well understood. To shed light on the topic, we scanned (fMRI) healthy adults during rest and during a task in which they discriminated pac-man configurations that formed or failed to form completed shapes (illusory and fragmented condition, respectively). Task activation differences (illusory-fragmented), resting-state functional connectivity, and multivariate patterns were identified on the cortical surface using 360 predefined parcels and 12 functional networks composed of such parcels. Brain activity flow mapping (ActFlow) was used to evaluate the likely involvement of resting-state connections for shape completion. We identified 36 differentially-active parcels including a posterior temporal region, PH, whose activity was consistent across 95% of observers. Significant task regions primarily occupied the secondary visual network but also incorporated the frontoparietal, dorsal attention, default mode, and cingulo-opercular networks. Each parcel's task activation difference could be modeled via its resting-state connections with the remaining parcels (r=.62, p<10-9), suggesting that such connections undergird shape completion. Functional connections from the dorsal attention network were key in modelling task activation differences in the secondary visual network. Dorsal attention and frontoparietal connections could also model activations in the remaining networks. Taken together, these results suggest that shape completion relies upon a sparsely distributed but densely interconnected network coalition that is centered in the secondary visual network, coordinated by the dorsal attention network, and inclusive of at least three other networks.

Tracking the completion of parts into whole objects: Retinotopic activation in response to illusory figures in the lateral occipital complex.

Illusory figures demonstrate the visual system's ability to integrate separate parts into coherent, whole objects. The present study was performed to track the neuronal object construction process in human observers, by incrementally manipulating the grouping strength within a given configuration until the emergence of a whole-object representation. Two tasks were employed: First, in the spatial localization task, object completion could facilitate performance and was task-relevant, whereas it was irrelevant in the second, luminance discrimination task. Concurrent functional magnetic resonance imaging (fMRI) used spatial localizers to locate brain regions representing task-critical illusory-figure parts to investigate whether the step-wise object construction process would modulate neural activity in these localized brain regions. The results revealed that both V1 and the lateral occipital complex (LOC, with sub-regions LO1 and LO2) were involved in Kanizsa figure processing. However, completion-specific activations were found predominantly in LOC, where neural activity exhibited a modulation in accord with the configuration's grouping strength, whether or not the configuration was relevant to performing the task at hand. Moreover, right LOC activations were confined to LO2 and responded primarily to surface and shape completions, whereas left LOC exhibited activations in both LO1 and LO2 and was related to encoding shape structures with more detail. Together, these results demonstrate that various grouping properties within a visual scene are integrated automatically in LOC, with sub-regions located in different hemispheres specializing in the component sub-processes that render completed objects.

Brain mechanisms for perceiving illusory lines in humans.

Illusory contours (ICs) are perceptions of visual borders despite absent contrast gradients. The psychophysical and neurobiological mechanisms of IC processes have been studied across species and diverse brain imaging/mapping techniques. Nonetheless, debate continues regarding whether IC sensitivity results from a (presumably) feedforward process within low-level visual cortices (V1/V2) or instead are processed first within higher-order brain regions, such as lateral occipital cortices (LOC). Studies in animal models, which generally favour a feedforward mechanism within V1/V2, have typically involved stimuli inducing IC lines. By contrast, studies in humans generally favour a mechanism where IC sensitivity is mediated by LOC and have typically involved stimuli inducing IC forms or shapes. Thus, the particular stimulus features used may strongly contribute to the model of IC sensitivity supported. To address this, we recorded visual evoked potentials (VEPs) while presenting human observers with an array of 10 inducers within the central 5°, two of which could be oriented to induce an IC line on a given trial. VEPs were analysed using an electrical neuroimaging framework. Sensitivity to the presence vs. absence of centrally-presented IC lines was first apparent at ∼200 ms post-stimulus onset and was evident as topographic differences across conditions. We also localized these differences to the LOC. The timing and localization of these effects are consistent with a model of IC sensitivity commencing within higher-level visual cortices. We propose that prior observations of effects within lower-tier cortices (V1/V2) are the result of feedback from IC sensitivity that originates instead within higher-tier cortices (LOC).

Infants Perceive Three-Dimensional Subjective Contours.

The addition of crossed horizontal disparity enhances the clarity of illusory contours compared to pictorial illusory contours and illusory contours with uncrossed horizontal disparity. Two infant-controlled habituation-dishabituation experiments explored the presence of this effect in infants 5 months of age. Experiment 1 examined whether infants are able to distinguish between a Kanizsa figure with crossed horizontal disparity and a Kanizsa figure with uncrossed horizontal disparity. Experiment 2 tested infants for their ability to differentiate between a Kanizsa figure with crossed horizontal disparity and a two-dimensional Kanizsa figure. The results provided evidence that the participants perceived the two- and the three-dimensional illusory Kanizsa contour, the illusory effect in which was strengthened by the addition of crossed horizontal disparity.

Cue-dependent circuits for illusory contours in humans.

Objects' borders are readily perceived despite absent contrast gradients, e.g. due to poor lighting or occlusion. In humans, a visual evoked potential (VEP) correlate of illusory contour (IC) sensitivity, the "IC effect", has been identified with an onset at ~90 ms and generators within bilateral lateral occipital cortices (LOC). The IC effect is observed across a wide range of stimulus parameters, though until now it always involved high-contrast achromatic stimuli. Whether IC perception and its brain mechanisms differ as a function of the type of stimulus cue remains unknown. Resolving such will provide insights on whether there is a unique or multiple solutions to how the brain binds together spatially fractionated information into a cohesive perception. Here, participants discriminated IC from no-contour (NC) control stimuli that were either comprised of low-contrast achromatic stimuli or instead isoluminant chromatic contrast stimuli (presumably biasing processing to the magnocellular and parvocellular pathways, respectively) on separate blocks of trials. Behavioural analyses revealed that ICs were readily perceived independently of the stimulus cue--i.e. when defined by either chromatic or luminance contrast. VEPs were analysed within an electrical neuroimaging framework and revealed a generally similar timing of IC effects across both stimulus contrasts (i.e. at ~90 ms). Additionally, an overall phase shift of the VEP on the order of ~30 ms was consistently observed in response to chromatic vs. luminance contrast independently of the presence/absence of ICs. Critically, topographic differences in the IC effect were observed over the ~110-160 ms period; different configurations of intracranial sources contributed to IC sensitivity as a function of stimulus contrast. Distributed source estimations localized these differences to LOC as well as V1/V2. The present data expand current models by demonstrating the existence of multiple, cue-dependent circuits in the brain for generating perceptions of illusory contours.

From local to global processing: the development of illusory contour perception.

Global visual processing is important for segmenting scenes, extracting form from background, and recognizing objects. Local processing involves attention to the local elements, contrast, and boundaries of an image at the expense of extracting a global percept. Previous work is inconclusive regarding the relative development of local and global processing. Some studies suggest that global perception is already present by 8 months of age, whereas others suggest that the ability arises during childhood and continues to develop during adolescence. We used a novel method to assess the development of global processing in 3- to 10-year-old children and an adult comparison group. We used Kanizsa illusory contours as an assay of global perception and measured responses on a touch-sensitive screen while monitoring eye position with a head-mounted eye tracker. Participants were tested using a similarity match-to-sample paradigm. Using converging measures, we found a clear developmental progression with age such that the youngest children performed near chance on the illusory contour discrimination, whereas 7- and 8-year-olds performed nearly perfectly, as did adults. There was clear evidence of a gradual shift from a local processing strategy to a global one; young children looked predominantly at and touched the "pacman" inducers of the illusory form, whereas older children and adults looked predominantly at and touched the middle of the form. These data show a prolonged developmental trajectory in appreciation of global form, with a transition from local to global visual processing between 4 and 7 years of age.

Visual search of illusory contours: The role of illusory contour clarity.

Illusory contours demonstrate an important function of the visual system-object inference from incomplete boundaries, which can arise from factors such as low luminance, camouflage, or occlusion. Illusory contours can be perceived with varying degrees of clarity depending on the features of their inducers. The present study aimed to evaluate whether illusory contour clarity influences visual search efficiency. Experiment 1 compared visual search performance for Kanizsa illusory stimuli and nonillusory inducer stimuli when manipulating inducer size as a clarity factor. Experiment 2 examined the effects of illusory contour clarity on visual search by manipulating the number of rings with missing arcs (i.e., line ends) comprising the inducers, for both illusory and nonillusory stimuli. To investigate whether surface alterations had an impact on visual search in Experiment 1, Experiment 3 examined search performance for Kanizsa-like stimuli formed from "smoothed" inducers compared with standard Kanizsa figures. The results of Experiments 1 and 2 indicated that while Kanizsa produced inefficient search, this was not contingent on the clarity of the illusory contours. Experiment 3 suggested that surface alterations of Kanizsa figures did impact visual search performance. Together, the results indicated that illusory contour clarity did not have much bearing on search performance. In certain conditions, Kanizsa figures even facilitated search compared with nonillusory stimuli, suggesting that rather than contour inference, surface features might have greater relevance in guiding visual attention.

Perception of illusory contours in children and adults: An eye-tracking study.

The eye-tracking study investigated the perception of subjective Kanizsa and Ehrenstein figures in adults and in children aged 3-4, 5-6, 7-8, and 9-11 years of age. More specifically, the distribution of looking at the inner stimulus part versus the inducing elements was measured for illusory figures, figures with real contours, and control displays. It was hypothesized that longer looking at the inner area of the illusory figures indicates global contour interpolation, whereas longer looking at the inducing elements indicates a local processing mode. According to the results, participants of all ages looked longer at the illusory Kanizsa and Ehrenstein contours than at the figures' inducing elements. However, performance was lowest in the children aged 3-4 years and increased during the preschool period. Moreover, the illusory contour displays elicited comparable visual responses as did the real contour displays. The use of the control displays that contained no contour information ensured that the participants' looking behavior was not driven by a spontaneous tendency to attend to the inner stimulus parts. The study confirms the view that sensitivity to illusory contours emerges very early in life.

Visual search of illusory contours: The role of illusory contour clarity.

Kanizsa-type illusory contours demonstrate an important function of the visual system-object inference from incomplete boundaries, which can be due to low luminance environments, camouflage, or occlusion. At a perceptual level, Kanizsa figures have been shown to have various degrees of clarity, depending on the features of the inducers. The aim of the present study is to evaluate whether contour clarity influences search efficiency of Kanizsa-type illusory contours. Experiment 1 will examine search for a Kanizsa-type illusory target among Kanizsa-type illusory distractors, by manipulating contour clarity using inducer size in three conditions, compared with search for a nonillusory perceptually grouped target among nonillusory perceptually grouped distractors with manipulated inducer size. Experiment 2 will address the effects of contour clarity on visual search by manipulating the number of arcs (i.e., line ends) comprising the inducers, in a visual search task of Kanizsa-type stimuli, compared with visual search for nonillusory grouped targets and distractors when the number of arcs are manipulated. To examine whether surface alterations had an impact on search in Experiment 1 due to changes in inducer size, Experiment 3 will examine search for Kanizsa stimuli formed from "smoothed" inducers, in comparison to search for Kanizsa stimuli used in Experiment 1. Together, these experiments will demonstrate whether contour clarity impacts visual search of illusory contours.

Visual saltation illusion induced by flashes of subjective contours.

The visual saltation illusion of a Kanizsa-type subjective triangle was demonstrated. After a subjective/real triangle was flashed twice at the same position, another subjective/real triangle was flashed at a displaced position. In a typical case, the second flash was perceived to occur midway between the first and third flash positions. This study showed that the rated illusion strength for the Kanizsa and real triangles largely depended on stimulus onset asynchrony and retinal eccentricity and that the illusion rating was the same between the Kanizsa and real gray triangles when they were presented on black disks (or inducers). When the real triangle was presented in isolation, the illusion rating was lower. Presenting flashes on disks appears to enhance the saltation illusion for both the Kanizsa and real triangles possibly due to a stronger crowding effect or shape changes of inducers enhancing the perception of object appearance and disappearance. Various types of saltation illusions with a Kanizsa triangle are demonstrated in a video.

From separate items to an integrated unit in visual working memory: Similarity chunking vs. configural grouping.

It is unclear whether sequentially presented items can be grouped into an ensemble by similarity chunking without the aid of the configural cue (e.g., Kanizsa figure). The current study systematically explored this issue by gradually weakening the configural cue until it was completely removed. The results showed that sequentially presented Pac-Men could be integrated into a Kanizsa figure for better memory only when they were successively presented in adjacent locations. When Pac-Men were presented at non-adjacent or unpredictable locations, they could be chunked by their orientation similarity (they all faced toward the fixation cross) to improve memory performance. Importantly, such orientation similarity alone was sufficient to evoke memory benefit, even when successively presented Pac-Men could not form a Kanizsa object structure. This memory benefit cannot simply be attributed to the statistic regularity or mental transformation based on the egocentric reference frame. However, the configural cue alone could not produce a memory benefit if these Pac-Men did not face toward the fixation cross. All of these results indicate that the memory benefit of sequentially presented items could occur by similarity chunking without the aid of the configural cue. The memory benefit based on the configural grouping was fragile when items were sequentially presented.

Number is not just an illusion: Discrete numerosity is encoded independently from perceived size.

While seminal theories suggest that nonsymbolic visual numerosity is mainly extracted from segmented items, more recent views advocate that numerosity cannot be processed independently of nonnumeric continuous features confounded with the numerical set (i.e., such as the density, the convex hull, etc.). To disentangle these accounts, here we employed two different visual illusions presented in isolation or in a merged condition (e.g., combining the effects of the two illusions). In particular, in a number comparison task, we concurrently manipulated both the perceived object segmentation by connecting items with Kanizsa-like illusory lines, and the perceived convex-hull/density of the set by embedding the stimuli in a Ponzo illusion context, keeping constant other low-level features. In Experiment 1, the two illusions were manipulated in a compatible direction (i.e., both triggering numerical underestimation), whereas in Experiment 2 they were manipulated in an incompatible direction (i.e., with the Ponzo illusion triggering numerical overestimation and the Kanizsa illusion numerical underestimation). Results from psychometric functions showed that, in the merged condition, the biases of each illusion summated (i.e., largest underestimation as compared with the conditions in which illusions were presented in isolation) in Experiment 1, while they averaged and competed against each other in Experiment 2. These findings suggest that discrete nonsymbolic numerosity can be extracted independently from continuous magnitudes. They also point to the need of more comprehensive theoretical views accounting for the operations by which both discrete elements and continuous variables are computed and integrated by the visual system.

Object-based grouping benefits without integrated feature representations in visual working memory.

Visual working memory (VWM) is typically considered to represent complete objects-that is, separate parts of an object are maintained as bound objects. Yet it remains unclear whether and how the features of disparate parts are integrated into a whole-object memory representation. Using a change detection paradigm, the present study investigated whether VWM performance varies as a function of grouping strength for features that either determine the grouped object (orientation) or that are not directly grouping relevant (color). Our results showed a large grouping benefit for grouping-relevant orientation features and, additionally, a much smaller, albeit reliable, benefit for grouping-irrelevant color features when both were potentially task relevant. By contrast, when color was the only task-relevant feature, no grouping benefit from the orientation feature was revealed both under lower or relatively high demands for precision. Together, these results indicate that different features of an object are stored independently in VWM; and an emerging, higher-order grouping structure does not automatically lead to an integrated representation of all available features of an object. Instead, an object benefit depends on the specific task demands, which may generate a linked, task-dependent representation of independent features.

Predictive coding feedback results in perceived illusory contours in a recurrent neural network.

Modern feedforward convolutional neural networks (CNNs) can now solve some computer vision tasks at super-human levels. However, these networks only roughly mimic human visual perception. One difference from human vision is that they do not appear to perceive illusory contours (e.g. Kanizsa squares) in the same way humans do. Physiological evidence from visual cortex suggests that the perception of illusory contours could involve feedback connections. Would recurrent feedback neural networks perceive illusory contours like humans? In this work we equip a deep feedforward convolutional network with brain-inspired recurrent dynamics. The network was first pretrained with an unsupervised reconstruction objective on a natural image dataset, to expose it to natural object contour statistics. Then, a classification decision head was added and the model was finetuned on a form discrimination task: squares vs. randomly oriented inducer shapes (no illusory contour). Finally, the model was tested with the unfamiliar "illusory contour" configuration: inducer shapes oriented to form an illusory square. Compared with feedforward baselines, the iterative "predictive coding" feedback resulted in more illusory contours being classified as physical squares. The perception of the illusory contour was measurable in the luminance profile of the image reconstructions produced by the model, demonstrating that the model really "sees" the illusion. Ablation studies revealed that natural image pretraining and feedback error correction are both critical to the perception of the illusion. Finally we validated our conclusions in a deeper network (VGG): adding the same predictive coding feedback dynamics again leads to the perception of illusory contours.

Kanizsa-figure object completion gates selection in the attentional blink.

Previous work has demonstrated that perceptual grouping modulates the selectivity of attention across space. By contrast, how grouping influences the allocation of attention over time is much less clear. This study investigated this issue, using an attentional blink (AB) paradigm to test how grouping influences the initial selection and the subsequent short-term memory consolidation of a target. On a given trial, two red Kanizsa-type targets (T1 and T2) with varying grouping strength were embedded in a rapid serial visual presentation stream of irrelevant distractors. Our results showed the typical AB finding: impaired identification of T2 when presented close in time following T1. Moreover, the AB was modulated by the T2 grouping-independently of the T1 structure-with stronger grouping leading to a decreased AB and overall higher performance. Conversely, a reversed pattern, namely an increased AB with increasing grouping strength was observed when the Kanizsa figure was not task-relevant. Together, these findings suggest that the grouping benefit emerges at early perceptual stages, automatically drawing attentional resources, thereby leading to either sustained benefits or transient costs-depending on the task-relevance of the grouped object. This indicates that grouping modulates processing of objects in time.

Disillusionment.

Recent research has shown that perceptual processes carry intrinsic affect. But prior studies have only manipulated the occurrence of perceptual processes by presenting two different stimulus categories. The present studies go beyond this by manipulating perceptual expectations for identical stimuli. Seven experiments demonstrated that objectively identical stimuli become visually disappointing and are liked less when they violate the expectation that an intrinsically pleasant perceptual process will occur compared to when there is no perceptual expectation. These effects were specific to violations of perceptual expectations. By using between-subjects designs, participants' insight into the experimental manipulation was prevented. In combination with the use of identical stimuli across conditions, this provides the most stringent test of the idea that perception is intrinsically (un-)pleasant yet. The results are related to predictive coding frameworks and provide an explanation for why people sometimes enjoy additional perceptual effort.

Global and local visual processing in autism: An objective assessment approach.

We examined global and local visual processing in autism spectrum disorder (ASD) via a match-to-sample task using Kanizsa illusory contours (KIC). School-aged children with ASD (n = 28) and age-matched typically developing controls (n = 22; 7-13 years) performed a sequential match-to-sample between a solid shape (sample) and two illusory alternatives. We tracked eye gaze and behavioral performance in two task conditions: one with and one without local interference from background noise elements. While analyses revealed lower accuracy and longer reaction time in ASD in the condition with local interference only, eye tracking robustly captured ASD-related global atypicalities across both conditions. Specifically, relative to controls, children with ASD showed decreased fixations to KIC centers, indicating reduced global perception. Notably, they did not differ from controls in regard to fixations to local elements or touch response location. These results indicate impaired global perception in the absence of heightened local processing in ASD. They also underscore the utility of eye-tracking measures as objective indices of global/local visual processing strategies in ASD. Autism Res 2017, 10: 1392-1404. © 2017 International Society for Autism Research, Wiley Periodicals, Inc.

The Kanizsa triangle illusion in foraging ants.

The Kanizsa triangle, wherein three Pac-Man configurations symmetrically face inwards, is a well-known illusion. By exposing foraging ants (Lasius niger) to Kanizsa-shaped honeydew solutions, we studied the origin of this illusion. More specifically, we examined whether foraging ants showed different movement reactions to local honeydew patterns formed by nestmates. This novel phenomenon could serve as an abstract model of the Kanizsa triangle illusion under the assumption that such an illusion could arise through the sum of each agent's limited global cognitions, because each agent could not perceive the entire subjective contours. Even a subjective consciousness consists of some parts which have no identical perception and could be an illusion. We succeeded in inducing foragers to move along the sides of a Kanizsa triangle when Pac-Man-shaped inducers were introduced. Furthermore, foragers appeared to form Y-shaped trajectories when dot-shaped or inverse Kanizsa inducers were used. Based on our findings, we propose an agent-based ant model that compares modelled behaviour with experimental phenomena. Our abstract model could be used to explain such cognitive phenomena for bottom-up processes, because ants cannot perceive the given subjective contours, instead simply move along the edges.