Cortical Surround Interactions and Perceptual Salience via Natural Scene Statistics

Spatial context in images induces perceptual phenomena associated with salience and modulates the responses of neurons in primary visual cortex (V1). However, the computational and ecological principles underlying contextual effects are incompletely understood. We introduce a model of natural images that includes grouping and segmentation of neighboring features based on their joint statistics, and we interpret the firing rates of V1 neurons as performing optimal recognition in this model. We show that this leads to a substantial generalization of divisive normalization, a computation that is ubiquitous in many neural areas and systems. A main novelty in our model is that the influence of the context on a target stimulus is determined by their degree of statistical dependence. We optimized the parameters of the model on natural image patches, and then simulated neural and perceptual responses on stimuli used in classical experiments. The model reproduces some rich and complex response patterns observed in V1, such as the contrast dependence, orientation tuning and spatial asymmetry of surround suppression, while also allowing for surround facilitation under conditions of weak stimulation. It also mimics the perceptual salience produced by simple displays, and leads to readily testable predictions. Our results provide a principled account of orientation-based contextual modulation in early vision and its sensitivity to the homogeneity and spatial arrangement of inputs, and lends statistical support to the theory that V1 computes visual salience.     

Temporal expectancy in the context of a theory of visual attention

Temporal expectation is expectation with respect to the timing of an event such as the appearance of a certain stimulus. In this paper, temporal expectancy is investigated in the context of the theory of visual attention (TVA), and we begin by summarizing the foundations of this theoretical framework. Next, we present a parametric experiment exploring the effects of temporal expectation on perceptual processing speed in cued single-stimulus letter recognition with unspeeded motor responses. The length of the cue–stimulus foreperiod was exponentially distributed with one of six hazard rates varying between blocks. We hypothesized that this manipulation would result in a distinct temporal expectation in each hazard rate condition. Stimulus exposures were varied such that both the temporal threshold of conscious perception (t₀ ms) and the perceptual processing speed (v letters s⁻¹) could be estimated using TVA. We found that the temporal threshold t₀ was unaffected by temporal expectation, but the perceptual processing speed v was a strikingly linear function of the logarithm of the hazard rate of the stimulus presentation. We argue that the effects on the v values were generated by changes in perceptual biases, suggesting that our perceptual biases are directly related to our temporal expectations.     

Analysis of spiking synchrony in visual cortex reveals distinct types of top-down modulation signals for spatial and object-based attention

The activity of a border ownership selective (BOS) neuron indicates where a foreground object is located relative to its (classical) receptive field (RF). A population of BOS neurons thus provides an important component of perceptual grouping, the organization of the visual scene into objects. In previous theoretical work, it has been suggested that this grouping mechanism is implemented by a population of dedicated grouping (“G”) cells that integrate the activity of the distributed feature cells representing an object and, by feedback, modulate the same cells, thus making them border ownership selective. The feedback modulation by G cells is thought to also provide the mechanism for object-based attention. A recent modeling study showed that modulatory common feedback, implemented by synapses with N-methyl-D-aspartate (NMDA)-type glutamate receptors, accounts for the experimentally observed synchrony in spike trains of BOS neurons and the shape of cross-correlations between them, including its dependence on the attentional state. However, that study was limited to pairs of BOS neurons with consistent border ownership preferences, defined as two neurons tuned to respond to the same visual object, in which attention decreases synchrony. But attention has also been shown to increase synchrony in neurons with inconsistent border ownership selectivity. Here we extend the computational model from the previous study to fully understand these effects of attention. We postulate the existence of a second type of G-cell that represents spatial attention by modulating the activity of all BOS cells in a spatially defined area. Simulations of this model show that a combination of spatial and object-based mechanisms fully accounts for the observed pattern of synchrony between BOS neurons. Our results suggest that modulatory feedback from G-cells may underlie both spatial and object-based attention.     

Perceptual Grouping Enhances Visual Plasticity

Visual perceptual learning, a manifestation of neural plasticity, refers to improvements in performance on a visual task achieved by training. Attention is known to play an important role in perceptual learning, given that the observer''s discriminative ability improves only for those stimulus feature that are attended. However, the distribution of attention can be severely constrained by perceptual grouping, a process whereby the visual system organizes the initial retinal input into candidate objects. Taken together, these two pieces of evidence suggest the interesting possibility that perceptual grouping might also affect perceptual learning, either directly or via attentional mechanisms. To address this issue, we conducted two experiments. During the training phase, participants attended to the contrast of the task-relevant stimulus (oriented grating), while two similar task-irrelevant stimuli were presented in the adjacent positions. One of the two flanking stimuli was perceptually grouped with the attended stimulus as a consequence of its similar orientation (Experiment 1) or because it was part of the same perceptual object (Experiment 2). A test phase followed the training phase at each location. Compared to the task-irrelevant no-grouping stimulus, orientation discrimination improved at the attended location. Critically, a perceptual learning effect equivalent to the one observed for the attended location also emerged for the task-irrelevant grouping stimulus, indicating that perceptual grouping induced a transfer of learning to the stimulus (or feature) being perceptually grouped with the task-relevant one. Our findings indicate that no voluntary effort to direct attention to the grouping stimulus or feature is necessary to enhance visual plasticity.     

Perceptual organization at attended and unattended locations

This study examined the effects of attention on forming perceptual units by proximity grouping and by uniform connectedness (UC). In Experiment 1 a row of three global letters defined by either proximity or UC was presented at the center of the visual field. Participants were asked to identify the letter in the middle of stimulus arrays while ignoring the flankers. The stimulus onset asynchrony (SOA) between stimulus arrays and masks varied between 180 and 500 ms. We found that responses to targets defined by proximity grouping were slower than to those defined by UC at median SOAs but there were no differences at short or long SOAs. Incongruent flankers slowed responses to targets and this flanker compatibility effect was larger for UC than for proximity-defined flankers. Experiment 2 examined the effects of spatial precueing on discrimination responses to proximity- and UC-defined targets. The advantage for targets defined by UC over targets defined by proximity grouping was greater at uncued relative to cued locations. The results suggest that the advantage for UC over proximity grouping in forming perceptual units is contingent on the stimuli not being fully attended, and that paying attention to the stimuli differentially benefits proximity grouping.     

Perceptual organization at attended and unattended locations

In order to perceive complex visual scenes, the human perceptual system has to organize discrete enti-ties in the visual field into chunks or perceptual units for higher-level processing. Perceptual organization is governed by Gestalt principles such as proximity, similarity, and continuity[1]. Thus spatially close ob-jects tend to be grouped together, as do elements that are similar to one another. Grouping based on the Ge-stalt laws (particularly proximity) is critical for the perception of…     

Components of Attention in Grapheme-Color Synesthesia: A Modeling Approach

Grapheme-color synesthesia is a condition where the perception of graphemes consistently and automatically evokes an experience of non-physical color. Many have studied how synesthesia affects the processing of achromatic graphemes, but less is known about the synesthetic processing of physically colored graphemes. Here, we investigated how the visual processing of colored letters is affected by the congruence or incongruence of synesthetic grapheme-color associations. We briefly presented graphemes (10–150 ms) to 9 grapheme-color synesthetes and to 9 control observers. Their task was to report as many letters (targets) as possible, while ignoring digit (distractors). Graphemes were either congruently or incongruently colored with the synesthetes’ reported grapheme-color association. A mathematical model, based on Bundesen’s (1990) Theory of Visual Attention (TVA), was fitted to each observer’s data, allowing us to estimate discrete components of visual attention. The models suggested that the synesthetes processed congruent letters faster than incongruent ones, and that they were able to retain more congruent letters in visual short-term memory, while the control group’s model parameters were not significantly affected by congruence. The increase in processing speed, when synesthetes process congruent letters, suggests that synesthesia affects the processing of letters at a perceptual level. To account for the benefit in processing speed, we propose that synesthetic associations become integrated into the categories of graphemes, and that letter colors are considered as evidence for making certain perceptual categorizations in the visual system. We also propose that enhanced visual short-term memory capacity for congruently colored graphemes can be explained by the synesthetes’ expertise regarding their specific grapheme-color associations.     

Perceptual grouping of biological motion promotes binocular rivalry

Investigation of perceptual rivalry between conflicting stimuli presented one to each eye can further understanding of the neural underpinnings of conscious visual perception. During rivalry, visual awareness fluctuates between perceptions of the two stimuli. Here, we demonstrate that high-level perceptual grouping can promote rivalry between stimulus pairs that would otherwise be perceived as nonrivalrous. Perceptual grouping was generated with point-light walker stimuli that simulate human motion, visible only as lights placed on the joints. Although such walking figures are unrecognizable when stationary, recognition judgments as complex as gender and identity can accurately be made from animated displays, demonstrating the efficiency with which our visual system can group dynamic local signals into a globally coherent walking figure. We find that point-light walker stimuli presented one to each eye and in different colors and configurations results in strong rivalry. However, rivalry is minimal when the two walkers are split between the eyes or both presented to one eye. This pattern of results suggests that processing animated walker figures promotes rivalry between signals from the two eyes rather than between higher-level representations of the walkers. This leads us to hypothesize that awareness during binocular rivalry involves the integrated activity of high-level perceptual mechanisms in conjunction with lower-level ocular suppression modulated via cortical feedback.     

Is Attention Based on Spatial Contextual Memory Preferentially Guided by Low Spatial Frequency Signals?

A popular model of visual perception states that coarse information (carried by low spatial frequencies) along the dorsal stream is rapidly transmitted to prefrontal and medial temporal areas, activating contextual information from memory, which can in turn constrain detailed input carried by high spatial frequencies arriving at a slower rate along the ventral visual stream, thus facilitating the processing of ambiguous visual stimuli. We were interested in testing whether this model contributes to memory-guided orienting of attention. In particular, we asked whether global, low-spatial frequency (LSF) inputs play a dominant role in triggering contextual memories in order to facilitate the processing of the upcoming target stimulus. We explored this question over four experiments. The first experiment replicated the LSF advantage reported in perceptual discrimination tasks by showing that participants were faster and more accurate at matching a low spatial frequency version of a scene, compared to a high spatial frequency version, to its original counterpart in a forced-choice task. The subsequent three experiments tested the relative contributions of low versus high spatial frequencies during memory-guided covert spatial attention orienting tasks. Replicating the effects of memory-guided attention, pre-exposure to scenes associated with specific spatial memories for target locations (memory cues) led to higher perceptual discrimination and faster response times to identify targets embedded in the scenes. However, either high or low spatial frequency cues were equally effective; LSF signals did not selectively or preferentially contribute to the memory-driven attention benefits to performance. Our results challenge a generalized model that LSFs activate contextual memories, which in turn bias attention and facilitate perception.     

Neuropsychological studies of object recognition

It is well established that disorders of visual perception are associated with lesions in the right hemisphere. Performances on tasks as disparate as the identification of objects from unusual views of objects drawn so as to overlap, of fragmented letters, of familiar faces, and of anomalous features in drawings, have been shown to be impaired in patients with focal right posterior lesions. A series of investigations are reviewed, directed towards analysing the basis of these deficits. Explanations in terms of primary visual impairment can be rejected, as can an account in terms of faulty figure-ground organization. It is argued that a wide variety of such perceptual deficits--all of which are concerned with meaningful visual stimuli--can be encompassed by the notion of faulty perceptual categorization at an early post-sensory stage of object recognition. Moreover, there is evidence suggesting that some of these various perceptual deficits can be mutually dissociated. The concept of perceptual categorization is discussed in the wider context of tentative model of object recognition.     

Top-Down Feedback in an HMAX-Like Cortical Model of Object Perception Based on Hierarchical Bayesian Networks and Belief Propagation

Hierarchical generative models, such as Bayesian networks, and belief propagation have been shown to provide a theoretical framework that can account for perceptual processes, including feedforward recognition and feedback modulation. The framework explains both psychophysical and physiological experimental data and maps well onto the hierarchical distributed cortical anatomy. However, the complexity required to model cortical processes makes inference, even using approximate methods, very computationally expensive. Thus, existing object perception models based on this approach are typically limited to tree-structured networks with no loops, use small toy examples or fail to account for certain perceptual aspects such as invariance to transformations or feedback reconstruction. In this study we develop a Bayesian network with an architecture similar to that of HMAX, a biologically-inspired hierarchical model of object recognition, and use loopy belief propagation to approximate the model operations (selectivity and invariance). Crucially, the resulting Bayesian network extends the functionality of HMAX by including top-down recursive feedback. Thus, the proposed model not only achieves successful feedforward recognition invariant to noise, occlusions, and changes in position and size, but is also able to reproduce modulatory effects such as illusory contour completion and attention. Our novel and rigorous methodology covers key aspects such as learning using a layerwise greedy algorithm, combining feedback information from multiple parents and reducing the number of operations required. Overall, this work extends an established model of object recognition to include high-level feedback modulation, based on state-of-the-art probabilistic approaches. The methodology employed, consistent with evidence from the visual cortex, can be potentially generalized to build models of hierarchical perceptual organization that include top-down and bottom-up interactions, for example, in other sensory modalities.     

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. Here, we explored the mechanism underlying the effects of these two modes of visual attention on the orientation selectivity of visual cortical neurons. To do this, we developed a layered microcircuit model. This model describes multiple orientation-specific microcircuits sharing their receptive fields and consisting of layers 2/3, 4, 5, and 6. These microcircuits represent a functional grouping of cortical neurons and mutually interact via lateral inhibition and excitatory connections between groups with similar selectivity. The individual microcircuits receive bottom-up visual stimuli and top-down attention in different layers. A crucial assumption of the model is that feature-based attention activates orientation-specific microcircuits for the relevant feature selectively, whereas spatial attention activates all microcircuits homogeneously, irrespective of their orientation selectivity. Consequently, our model simultaneously accounts for the multiplicative scaling of neuronal responses in spatial attention and the additive modulations of orientation tuning curves in feature-based attention, which have been observed widely in various visual cortical areas. Simulations of the model predict contrasting differences between excitatory and inhibitory neurons in the two modes of attentional modulations. Furthermore, the model replicates the modulation of the psychophysical discriminability of visual stimuli in the presence of external noise. Our layered model with a biologically suggested laminar structure describes the basic circuit mechanism underlying the attention-mode specific modulations of neuronal responses and visual perception.     

On the role of medial geometry in human vision.

A key challenge underlying theories of vision is how the spatially restricted, retinotopically represented feature analysis can be integrated to form abstract, coordinate-free object models. A resolution likely depends on the use of intermediate-level representations which can on the one hand be populated by local features and on the other hand be used as atomic units underlying the formation of, and interaction with, object hypotheses. The precise structure of this intermediate representation derives from the varied requirements of a range of visual tasks which motivate a significant role for incorporating a geometry of visual form. The need to integrate input from features capturing surface properties such as texture, shading, motion, color, etc., as well as from features capturing surface discontinuities such as silhouettes, T-junctions, etc., implies a geometry which captures both regional and boundary aspects. Curves, as a geometric model of boundaries, have been extensively used as an intermediate representation in computational, perceptual, and physiological studies, while the use of the medial axis (MA) has been popular mainly in computer vision as a geometric region-based model of the interior of closed boundaries. We extend the traditional model of the MA to represent images, where each MA segment represents a region of the image which we call a visual fragment. We present a unified theory of perceptual grouping and object recognition where through various sequences of transformations of the MA representation, visual fragments are grouped in various configurations to form object hypotheses, and are related to stored models. The mechanisms underlying both the computation and the transformation of the MA is a lateral wave propagation model. Recent psychophysical experiments depicting contrast sensitivity map peaks at the medial axes of stimuli, and experiments on perceptual filling-in, and brightness induction and modulation, are consistent with both the use of an MA representation and a propagation-based scheme. Also, recent neurophysiological recordings in V1 correlate with the MA hypothesis and a horizontal propagation scheme. This evidence supports a geometric computational paradigm for processing sensory data where both dynamic in-plane propagation and feedforward-feedback connections play an integral role.     

Processing local signals into global patterns

Perceptual organization or grouping is one of the central issues in vision research. Recent reports in the neuroimaging literature suggest that perceptual organization is mediated by distributed visual areas that range from the primary visual cortex (V1) to higher visual areas, depending on the availability of grouping cues and on the weight of contribution of each visual area. Evidence suggests that grouping by proximity and collinearity, and also perhaps filling-in, involve V1, whereas grouping by similarity and symmetry seems to depend on activation of higher visual areas. Further studies should include deliberate controls for confounding factors such as attentional artifacts and radial orientation bias, to clarify how spatiotemporal information in visual areas is integrated to give rise to perceptual organization.     

Space-variant visual processing: spatially limited visual channels

A multichannel model incorporating visual inhomogeneity is presented in this paper. The parameters that describe inhomogeneity have been experimentally obtained both at threshold and in several suprathreshold conditions. At threshold, probability summation is taken into account in order to determine the spatial extent of visual channels from experimental data showing an asymptotic increase in sensitivity with increasing grating area. At suprathreshold contrast, the region where luminance variations at several scales are visible has also been found. The results support a spatially limited multichannel model of early visual processing and set out a basis for studying perceptual phenomena from the viewpoint of linear space-variant visual processing.     

The Competitive Influences of Perceptual Load and Working Memory Guidance on Selective Attention

The perceptual load theory in selective attention literature proposes that the interference from task-irrelevant distractor is eliminated when perceptual capacity is fully consumed by task-relevant information. However, the biased competition model suggests that the contents of working memory (WM) can guide attentional selection automatically, even when this guidance is detrimental to visual search. An intriguing but unsolved question is what will happen when selective attention is influenced by both perceptual load and WM guidance. To study this issue, behavioral performances and event-related potentials (ERPs) were recorded when participants were presented with a cue to either identify or hold in memory and had to perform a visual search task subsequently, under conditions of low or high perceptual load. Behavioural data showed that high perceptual load eliminated the attentional capture by WM. The ERP results revealed an obvious WM guidance effect in P1 component with invalid trials eliciting larger P1 than neutral trials, regardless of the level of perceptual load. The interaction between perceptual load and WM guidance was significant for the posterior N1 component. The memory guidance effect on N1 was eliminated by high perceptual load. Standardized Low Resolution Electrical Tomography Analysis (sLORETA) showed that the WM guidance effect and the perceptual load effect on attention can be localized into the occipital area and parietal lobe, respectively. Merely identifying the cue produced no effect on the P1 or N1 component. These results suggest that in selective attention, the information held in WM could capture attention at the early stage of visual processing in the occipital cortex. Interestingly, this initial capture of attention by WM could be modulated by the level of perceptual load and the parietal lobe mediates target selection at the discrimination stage.     

Effects of lipid rafts on dynamics of retroviral entry and trafficking: Quantitative analysis

The association of cell surface receptors with sterol-sphingolipid-enriched microdomains of the plasma membrane, so-called lipid rafts, may affect the receptor-mediated entry and trafficking dynamics of viruses. A model retrovirus, subgroup A avian sarcoma and leukosis virus (ASLV-A), can initiate infection by binding to either of two forms of the tumor virus subgroup A (TVA) receptor, a lipid-raft-associated glycosylphosphatidylinositol (GPI)-anchored receptor (TVA800) or a transmembrane receptor (TVA950). Narayan et al. previously found that virus particles bound to TVA950 were more rapidly internalized than virions bound to TVA800, and the internalization via TVA950 exhibited biphasic kinetics. To explore potential molecular mechanisms for these results we developed a mathematical model that accounts for internalization of viruses through cellular pits, trafficking to an endosomal compartment where fusion occurs, and viral DNA synthesis. By fitting the model to experimental data we found that viruses bound to TVA950 were internalized up to 2.6-fold more rapidly than viruses bound to TVA800. Two- to threefold greater lateral diffusivities of transmembrane proteins, relative to GPI-anchored proteins, observed in other systems, suggest that the internalization rate of ASLV-A is diffusion-limited. Furthermore, by allowing for recycling of internalized TVA950-bound viruses back to the cell surface, we can account for the observed biphasic internalization kinetics. This mechanism is also consistent with the observed slower rate of DNA synthesis for viruses that enter via TVA950. Overall, the model provides a means to generate new experimentally testable hypotheses and sets a foundation for building a quantitative and integrated understanding of viral entry, trafficking, and intracellular dynamics.     

Visual perception of texture regularity: Conjoint measurements and a wavelet response-distribution model

Texture regularity, such as the repeating pattern in a carpet, brickwork or tree bark, is a ubiquitous feature of the visual world. The perception of regularity has generally been studied using multi-element textures in which the degree of regularity has been manipulated by adding random jitter to the elements’ positions. Here we used three-factor Maximum Likelihood Conjoint Measurement (MLCM) for the first time to investigate the encoding of regularity information under more complex conditions in which element spacing and size, in addition to positional jitter, were manipulated. Human observers were presented with large numbers of pairs of multi-element stimuli with varying levels of the three factors, and indicated on each trial which stimulus appeared more regular. All three factors contributed to regularity perception. Jitter, as expected, strongly affected regularity perception. This effect of jitter on regularity perception is strongest at small element spacing and large texture element size, suggesting that the visual system utilizes the edge-to-edge distance between elements as the basis for regularity judgments. We then examined how the responses of a bank of Gabor wavelet spatial filters might account for our results. Our analysis indicates that the peakedness of the spatial frequency (SF) distribution, a previously favored proposal, is insufficient for regularity encoding since it varied more with element spacing and size than with jitter. Instead, our results support the idea that the visual system may extract texture regularity information from the moments of the SF-distribution across orientation. In our best-performing model, the variance of SF-distribution skew across orientations can explain 70% of the variance of estimated texture regularity from our data, suggesting that it could provide a candidate read-out for perceived regularity.     

The peri-saccadic perception of objects and space

Eye movements affect object localization and object recognition. Around saccade onset, briefly flashed stimuli appear compressed towards the saccade target, receptive fields dynamically change position, and the recognition of objects near the saccade target is improved. These effects have been attributed to different mechanisms. We provide a unifying account of peri-saccadic perception explaining all three phenomena by a quantitative computational approach simulating cortical cell responses on the population level. Contrary to the common view of spatial attention as a spotlight, our model suggests that oculomotor feedback alters the receptive field structure in multiple visual areas at an intermediate level of the cortical hierarchy to dynamically recruit cells for processing a relevant part of the visual field. The compression of visual space occurs at the expense of this locally enhanced processing capacity.