Evolution of the Wilson-Cowan equations.

The Wilson-Cowan equations were developed to provide a simplified yet powerful description of neural network dynamics. As such, they embraced nonlinear dynamics, but in an interpretable form. Most importantly, it was the first mathematical formulation to emphasize the significance of interactions between excitatory and inhibitory neural populations, thereby incorporating both cooperation and competition. Subsequent research by many has documented the Wilson-Cowan significance in such diverse fields as visual hallucinations, memory, binocular rivalry, and epilepsy. The fact that these equations are still being used to elucidate a wide range of phenomena attests to their validity as a dynamical approximation to more detailed descriptions of complex neural computations.

Hyperchaos in Wilson-Cowan oscillator circuits.

The Wilson-Cowan equations were originally shown to produce limit cycle oscillations for a range of parameters. Others subsequently showed that two coupled Wilson-Cowan oscillators could produce chaos, especially if the oscillator coupling was from inhibitory interneurons of one oscillator to excitatory neurons of the other. Here this is extended to show that chains, grids, and sparse networks of Wilson-Cowan oscillators generate hyperchaos with linearly increasing complexity as the number of oscillators increases. As there is now evidence that humans can voluntarily generate hyperchaotic visuomotor sequences, these results are particularly relevant to the unpredictability of a range of human behaviors. These also include incipient senescence in aging, effects of concussive brain injuries, autism, and perhaps also intelligence and creativity.NEW & NOTEWORTHY This paper represents an exploration of hyperchaos in coupled Wilson-Cowan equations. Results show that hyperchaos (number of positive Lyapunov exponents) grows linearly with the number of oscillators in the array and leads to high levels of unpredictability in the neural response.

Voluntary Generation of Hyperchaotic Visuo-Motor Patterns.

Unpredictable escape behaviour is an integral part of the repertoire of many prey species. Many social species, including humans, also employ unpredictability to enhance their chances of success in social interactions with conspecifics. However, it is unclear what mechanisms provide the behavioural and neural bases of this unpredictability, which might result either from randomness or from chaotic dynamics. A novel paradigm described here demonstrates that unpredictable behaviour generated voluntarily in a simple visuomotor task represents high dimensional chaos, or hyperchaos, but not randomness. The exponential decay of predictability was also shown to be longer in older adults. As chaos has been observed among cortical neurons, this provides a plausible neural basis for hyperchaos. These results thus provide evidence that voluntary unpredictable behaviour can result from neural hyperchaos. This work has implications for the study of autism, aftereffects of concussions, early dementia, and the concept of free will.

Binocular contrast, stereopsis, and rivalry: Toward a dynamical synthesis.

It is well known that small orientation differences between two monocular gratings fuse to generate a stereoscopic perception of tilt, while large differences trigger binocular rivalry. In addition, unequal monocular contrasts combine nonlinearly to generate binocular contrast. A nonlinear neural model is developed here to account for binocular contrast, fusion at small orientation differences, and rivalry at large differences. The model also accounts for hysteresis in the transition between fusion and rivalry. Finally, the model predicts that interocular contrast differences between fusible gratings will produce a reduced tilt percept, and experiments reported here support this. Key to the model is the presence of two classes of inhibitory interneurons: one operating on similar orientations to normalize interocular contrast (IN), and one operating across large orientation differences to generate rivalry (IR). Critically, the IN neurons switch off the IR neurons driven by the other eye, thus permitting fusion of binocular plaids.

Face perception develops similarly across viewpoint in children and adolescents with and without autism spectrum disorder.

Atypical face perception has been associated with the socio-communicative difficulties that characterize autism spectrum disorder (ASD). Growing evidence, however, suggests that a widespread impairment in face perception is not as common as once thought. One important issue arising with the interpretation of this literature is the relationship between face processing and a more general perceptual tendency to focus on local rather than global information. Previous work has demonstrated that when discriminating faces presented from the same view, older adolescents and adults with ASD perform similarly to typically developing individuals. When faces are presented from different views, however, they perform more poorly-specifically, when access to local cues is minimized. In this study, we assessed the cross-sectional development of face identity discrimination across viewpoint using same- and different-view conditions in children and adolescents with and without ASD. Contrary to the findings in adults, our results revealed that all participants experienced greater difficulty identifying faces from different views than from same views, and demonstrated similar age-expected improvements in performance across tasks. These results suggest that differences in face discrimination across views may only emerge beyond the age of 15 years in ASD.

Position selectivity in face-sensitive visual cortex to facial and nonfacial stimuli: an fMRI study.

BACKGROUND: Evidence for position sensitivity in object-selective visual areas has been building. On one hand, most of the relevant studies have utilized stimuli for which the areas are optimally selective and examine small sections of cortex. On the other hand, visual field maps established with nonspecific stimuli have been found in increasingly large areas of visual cortex, though generally not in areas primarily responsive to faces. METHODS: fMRI was used to study the position sensitivity of the occipital face area (OFA) and the fusiform face area (FFA) to both standard rotating wedge retinotopic mapping stimuli and quadrant presentations of synthetic facial stimuli. Analysis methods utilized were both typical, that is, mean univariate BOLD signals and multivoxel pattern analysis (MVPA), and novel, that is, distribution of voxels to pattern classifiers and use of responses to nonfacial retinotopic mapping stimuli to classify responses to facial stimuli. RESULTS: Polar angle sensitivity was exhibited to standard retinotopic mapping stimuli with a stronger contralateral bias in OFA than in FFA, a stronger bias toward the vertical meridian in FFA than in OFA, and a bias across both areas toward the inferior visual field. Contralateral hemispheric lateralization of both areas was again shown using synthetic face stimuli based on univariate BOLD signals, MVPA, and the biased contribution of voxels toward multivariate classifiers discriminating the contralateral visual field. Classifiers based on polar angle responsivity were used to classify the patterns of activation above chance levels to face stimuli in the OFA but not in the FFA. CONCLUSIONS: Both the OFA and FFA exhibit quadrant sensitivity to face stimuli, though the OFA exhibits greater position responsivity across stimuli than the FFA and includes overlap in the response pattern to the disparate stimulus types. Such biases are consistent with varying position sensitivity along different surfaces of occipito-temporal cortex.

A closer look at four-dot masking of a foveated target.

Four-dot masking with a common onset mask was recently demonstrated in a fully attended and foveated target (Filmer, Mattingley & Dux, 2015). Here, we replicate and extend this finding by directly comparing a four-dot mask with an annulus mask while probing masking as a function of mask duration, and target-mask separation. Our results suggest that while an annulus mask operates via spatially local contour interactions, a four-dot mask operates through spatially global mechanisms. We also measure how the visual system's representation of an oriented bar is impacted by a four-dot mask, and find that masking here does not degrade the precision of perceived targets, but instead appears to be driven exclusively by rendering the target completely invisible.

Detection of periodic motion trajectories: Effects of frequency and radius.

Periodic trajectories are an important component of biological motion. Or, Thabet, Wilkinson, and Wilson (2011) studied radial frequency (RF) motion trajectory detection and concluded that, for RF2-5 trajectories, the threshold function paralleled that of static RF patterns. We have extended Or et al.'s (2011) findings to a broader range of RFs (three to 24 cycles) and across a 4-fold range of radii (1°-4°). We report that (a) thresholds for RF trajectories decrease as a power function of RF for low RF trajectories (three to six cycles) before approaching an asymptote at high RFs (12-24 cycles); (b) detection thresholds for RF trajectories scale proportionally with radius; and (c) there is no lower versus upper field advantage in the parafoveal field for stimuli displaced from fixation on the vertical midline. The results are compared to earlier findings for static RF thresholds, and we argue that our findings support the existence of parallel spatial and temporal processing channels that may contribute to both action perception and production.

Effect of motion discontinuities on discrimination of periodic trajectories.

Many biologically important motions are described by periodic trajectories. Radial frequency (RF) trajectories are one example, in which the motion of a difference of Gaussians (DOG) target moves along a path described by a sinusoidal deviation of the radius from a perfect circle (Or, Thabet, Wilkinson, & Wilson, 2011). Here we explore the hypothesis that visual processing of RF trajectories involves global spatio-temporal processes that are disrupted by motion discontinuity. To test this hypothesis, RF trajectories were used that interspersed smooth, continuous motion with three or four discontinuous jumps to other portions of the trajectory. These jumps were arranged so that the entire trajectory was traversed in the same amount of time as in the continuous motion control condition. The motion discontinuities increased thresholds by a factor of approximately 2.1 relative to continuous motion. This result provides support for global spatio-temporal processing of RF motion trajectories. Comparison with previous results suggests that motion discontinuities erase memory for earlier parts of the trajectory, thereby causing thresholds to be based on only the final segment viewed. Finally, it is shown that RF trajectories obey the 1/3 power law characteristic of biological motion.

The Caledonian face test: A new test of face discrimination.

This study aimed to develop a clinical test of face perception which is applicable to a wide range of patients and can capture normal variability. The Caledonian face test utilises synthetic faces which combine simplicity with sufficient realism to permit individual identification. Face discrimination thresholds (i.e. minimum difference between faces required for accurate discrimination) were determined in an "odd-one-out" task. The difference between faces was controlled by an adaptive QUEST procedure. A broad range of face discrimination sensitivity was determined from a group (N=52) of young adults (mean 5.75%; SD 1.18; range 3.33-8.84%). The test is fast (3-4 min), repeatable (test-re-test r(2)=0.795) and demonstrates a significant inversion effect. The potential to identify impairments of face discrimination was evaluated by testing LM who reported a lifelong difficulty with face perception. While LM's impairment for two established face tests was close to the criterion for significance (Z-scores of -2.20 and -2.27) for the Caledonian face test, her Z-score was -7.26, implying a more than threefold higher sensitivity. The new face test provides a quantifiable and repeatable assessment of face discrimination ability. The enhanced sensitivity suggests that the Caledonian face test may be capable of detecting more subtle impairments of face perception than available tests.

Masking with faces in central visual field under a variety of temporal schedules.

With a few exceptions, previous studies have explored masking using either a backward mask or a common onset trailing mask, but not both. In a series of experiments, we demonstrate the use of faces in central visual field as a viable method to study the relationship between these two types of mask schedule. We tested observers in a two alternative forced choice face identification task, where both target and mask comprised synthetic faces, and show that a simple model can successfully predict masking across a variety of masking schedules ranging from a backward mask to a common onset trailing mask and a number of intermediate variations. Our data are well accounted for by a window of sensitivity to mask interference that is centered at around 100 ms.

The PCA learning effect: An emerging correlate of face memory during childhood.

Human adults implicitly learn the prototype and the principal components of the variability distinguishing faces (Gao & Wilson, 2014). Here we measured the implicit learning effect in adults and 9-year-olds, and with a modified child-friendly procedure, in 7-year-olds. All age groups showed the implicit learning effect by falsely recognizing the average (the prototype effect) and the principal component faces as having been seen (the PCA learning effect). The PCA learning effect, but not the prototype effect increased between 9years of age and adulthood and at both ages was the better predictor of memory for the actually studied faces. In contrast, for the 7-year-olds, the better predictor of face memory was the prototype effect. The pattern suggests that there may be a developmental change between ages 7 and 9 in the mechanism underlying memory for faces. We provide the first evidence that children as young as age 7 can extract the most important dimensions of variation represented by principal components among individual faces, a key ability that grows stronger with age and comes to underlie memory for faces.

From orientations to objects: Configural processing in the ventral stream.

The ventral or form vision hierarchy comprises a sequence of cortical areas in which successively more complex visual attributes are extracted, beginning with contour orientations in V1 and culminating in face and object representations at the highest levels. In addition, ventral areas exhibit increasing receptive field diameter by a factor of approximately three from area to area, and conversely neuron density decreases. We argue here that this is consistent with configural combination of adjacent orientations to form curves or angles, followed by combination of these to form descriptions of object shapes. Substantial data from psychophysics, functional magnetic resonance imaging (fMRI), and neurophysiology support this organization, and computational models consistent with it have also been proposed. We further argue that a key to the role of the ventral stream is dimensionality reduction in object representations.

Detection and recognition of angular frequency patterns.

Previous research has extensively explored visual encoding of smoothly curved, closed contours described by sinusoidal variation of pattern radius as a function of polar angle (RF patterns). Although the contours of many biologically significant objects are curved, we also confront shapes with a more jagged and angular appearance. To study these, we introduce here a novel class of visual stimuli that deform smoothly from a circle to an equilateral polygon with N sides (AF patterns). Threshold measurements reveal that both AF and RF patterns can be discriminated from circles at the same deformation amplitude, approximately 18.0arcsec, which is in the hyperacuity range. Thresholds were slightly higher for patterns with 3.0 cycles than for those with 5.0 cycles. Discrimination between AF and RF patterns was 75% correct at an amplitude that was approximately 3.0 times the threshold amplitude, which implies that AF and RF patterns activate different neural populations. Experiments with jittered patterns in which the contour was broken into several pieces and shifted inward or outward had much less effect on AF patterns than on RF patterns. Similarly, thresholds for single angles of AF patterns showed no significant difference from thresholds for the entire AF pattern. Taken together, these results imply that the visual system incorporates angles explicitly in the representation of closed object contours, but it suggests that angular contours are represented more locally than are curved contours.

Atypical Face Perception in Autism: A Point of View?

Face perception is the most commonly used visual metric of social perception in autism. However, when found to be atypical, the origin of face perception differences in autism is contentious. One hypothesis proposes that a locally oriented visual analysis, characteristic of individuals with autism, ultimately affects performance on face tasks where a global analysis is optimal. The objective of this study was to evaluate this hypothesis by assessing face identity discrimination with synthetic faces presented with and without changes in viewpoint, with the former condition minimizing access to local face attributes used for identity discrimination. Twenty-eight individuals with autism and 30 neurotypical participants performed a face identity discrimination task. Stimuli were synthetic faces extracted from traditional face photographs in both front and 20° side viewpoints, digitized from 37 points to provide a continuous measure of facial geometry. Face identity discrimination thresholds were obtained using a two-alternative, temporal forced choice match-to-sample paradigm. Analyses revealed an interaction between group and condition, with group differences found only for the viewpoint change condition, where performance in the autism group was decreased compared to that of neurotypical participants. The selective decrease in performance for the viewpoint change condition suggests that face identity discrimination in autism is more difficult when access to local cues is minimized, and/or when dependence on integrative analysis is increased. These results lend support to a perceptual contribution of atypical face perception in autism.

Bandwidths for the perception of head orientation decrease during childhood.

Adults use the orientation of people's heads as a cue to the focus of their attention. We examined developmental changes in mechanisms underlying sensitivity to head orientation during childhood. Eight-, 10-, 12-year-olds, and adults were adapted to a frontal face view or a 20° left or right side view before judging the orientation of a face at or near frontal. After frontal adaptation, there were no age differences in judgments of head orientation. However, after adaptation to a 20° left or right side view, aftereffects were larger and sensitivity to head orientation was lower in 8- and 10-year-olds than in adults, with no difference between 12-year-olds and adults. A computational model indicates that these results can be modeled as a consequence of decreasing neural tuning bandwidths and decreasing additive internal noise during childhood, and/or as a consequence of increasing inhibition during childhood. These results provide the first evidence that neural mechanisms underlying sensitivity to head orientation undergo considerable refinement during childhood.

Neural correlates of radial frequency trajectory perception in the human brain.

Radial frequency (RF) motion trajectories are visual stimuli that consist of a difference of Gaussians moving along a closed trajectory defined by a sinusoidal variation of the radius relative to a circular path. In the current study, multivoxel fMRI analyses demonstrated that spatial patterns of activity in visual regions V2, V3, and MT can predict RF motion trajectory shape regardless of whether an observer can behaviorally identify the shape or not. This result suggests that processing in these regions is concerned with local properties of the trajectories and not directly linked with a conscious percept of global trajectory shape. Whole-brain analyses show that RF motion trajectories also evoke premotor and posterior parietal cortical activity that may be a neural correlate of shape recognizability. Further, comparisons with activity evoked by static versions of the RF shapes reveal cue-invariant processing in regions of the posterior parietal and occipitotemporal cortices. Interestingly, the RF motion trajectories evoke patterns of dorsal visual stream cortical activity typical of visually guided movement preparation or action observation, suggesting that these stimuli may be processed as potential motor actions rather than as purely visual experiences.

Implicit learning of geometric eigenfaces.

The human visual system can implicitly extract a prototype of encountered visual objects (Posner & Keele, 1968). While learning a prototype provides an efficient way of encoding objects at the category level, discrimination among individual objects requires encoding of variations among them as well. Here we show that in addition to the prototype, human adults also implicitly learn the feature correlations that capture the most significant geometric variations among faces. After studying a group of synthetic faces, observers mistook as seen previously unseen faces representing the first two principal components (eigenfaces, Turk & Pentland, 1991) of the studied faces at significantly higher rates than the correct recognition of the faces actually studied. Implicit learning of the most significant eigenfaces provides an optimal way for encoding variations among faces. The data thus extend the types of summary statistics that can be implicitly extracted by the visual system to include several principal components.

The neural representation of face space dimensions.

Functional neural imaging studies have identified a network of brain areas that are more active to faces than to other objects. However, it remains largely unclear how these areas encode individual facial identity. To investigate the neural representations of facial identity, we constructed a multidimensional face space structure, whose dimensions were derived from geometric information of faces using the Principal Component Analysis (PCA). Using fMRI, we recorded participants' neural responses when viewing blocks of faces that differed only on one dimension within a block. Although the response magnitudes to different blocks of faces did not differ in a univariate analysis, multi-voxel pattern analysis revealed distinct patterns related to different face space dimensions in brain areas that have a higher response magnitude to faces than to other objects. The results indicate that dimensions of the face space are encoded in the face-selective brain areas in a spatially distributed way.

Implicit face prototype learning from geometric information.

There is evidence that humans implicitly learn an average or prototype of previously studied faces, as the unseen face prototype is falsely recognized as having been learned (Solso & McCarthy, 1981). Here we investigated the extent and nature of face prototype formation where observers' memory was tested after they studied synthetic faces defined purely in geometric terms in a multidimensional face space. We found a strong prototype effect: The basic results showed that the unseen prototype averaged from the studied faces was falsely identified as learned at a rate of 86.3%, whereas individual studied faces were identified correctly 66.3% of the time and the distractors were incorrectly identified as having been learned only 32.4% of the time. This prototype learning lasted at least 1 week. Face prototype learning occurred even when the studied faces were further from the unseen prototype than the median variation in the population. Prototype memory formation was evident in addition to memory formation of studied face exemplars as demonstrated in our models. Additional studies showed that the prototype effect can be generalized across viewpoints, and head shape and internal features separately contribute to prototype formation. Thus, implicit face prototype extraction in a multidimensional space is a very general aspect of geometric face learning.