Stages Of Cortical Visual Processing Research Articles

Neural Correlates of Crowding in Macaque Area V4.

Visual crowding refers to the phenomenon where a target object that is easily identifiable in isolation becomes difficult to recognize when surrounded by other stimuli (distractors). Many psychophysical studies have investigated this phenomenon and proposed alternative models for the underlying mechanisms. One prominent hypothesis, albeit with mixed psychophysical support, posits that crowding arises from the loss of information due to pooled encoding of features from target and distractor stimuli in the early stages of cortical visual processing. However, neurophysiological studies have not rigorously tested this hypothesis. We studied the responses of single neurons in macaque (one male, one female) area V4, an intermediate stage of the object-processing pathway, to parametrically designed crowded displays and texture statistics-matched metameric counterparts. Our investigations reveal striking parallels between how crowding parameters-number, distance, and position of distractors-influence human psychophysical performance and V4 shape selectivity. Importantly, we also found that enhancing the salience of a target stimulus could alleviate crowding effects in highly cluttered scenes, and this could be temporally protracted reflecting a dynamical process. Thus, a pooled encoding of nearby stimuli cannot explain the observed responses, and we propose an alternative model where V4 neurons preferentially encode salient stimuli in crowded displays. Overall, we conclude that the magnitude of crowding effects is determined not just by the number of distractors and target-distractor separation but also by the relative salience of targets versus distractors based on their feature attributes-the similarity of distractors and the contrast between target and distractor stimuli.

Neurophysiology: The Three-Dimensional Building Blocks of Object Vision

A new study of the macaque visual cortex has revealed that visual area V4 performs substantial analysis of solid shape structure. The findings draw new attention to the embedding of local three-dimensional shape analysis into the early cortical stages of visual processing.

Task-related activity in human visual cortex.

The brain exhibits widespread endogenous responses in the absence of visual stimuli, even at the earliest stages of visual cortical processing. Such responses have been studied in monkeys using optical imaging with a limited field of view over visual cortex. Here, we used functional MRI (fMRI) in human participants to study the link between arousal and endogenous responses in visual cortex. The response that we observed was tightly entrained to task timing, was spatially extensive, and was independent of visual stimulation. We found that this response follows dynamics similar to that of pupil size and heart rate, suggesting that task-related activity is related to arousal. Finally, we found that higher reward increased response amplitude while decreasing its trial-to-trial variability (i.e., the noise). Computational simulations suggest that increased temporal precision underlies both of these observations. Our findings are consistent with optical imaging studies in monkeys and support the notion that arousal increases precision of neural activity.

Representational untangling by the firing rate nonlinearity in V1 simple cells.

An important computational goal of the visual system is 'representational untangling' (RU): representing increasingly complex features of visual scenes in an easily decodable format. RU is typically assumed to be achieved in high-level visual cortices via several stages of cortical processing. Here we show, using a canonical population coding model, that RU of low-level orientation information is already performed at the first cortical stage of visual processing, but not before that, by a fundamental cellular-level property: the thresholded firing rate nonlinearity of simple cells in the primary visual cortex (V1). We identified specific, experimentally measurable parameters that determined the optimal firing threshold for RU and found that the thresholds of V1 simple cells extracted from in vivo recordings in awake behaving mice were near optimal. These results suggest that information re-formatting, rather than maximisation, may already be a relevant computational goal for the early visual system.

Automatic Detection of Orientation Contrast Occurs at Early but Not Earliest Stages of Visual Cortical Processing in Humans.

Orientation contrast is formed when some elements orient differently from their surroundings. Although orientation contrast can be processed in the absence of top-down attention, the underlying neural mechanism for this automatic processing in humans is controversial. In particular, whether automatic detection of orientation contrast occurs at the initial feedforward stage in the primary visual cortex (i.e., V1) remains unclear. Here, we used event-related potentials (ERPs) to examine the automatic processing of orientation contrast in humans. In three experiments, participants completed a task at fixation while orientation contrasts were presented in the periphery, either in the upper visual field (UVF) or the lower visual field (LVF). All experiments showed significant positive potentials evoked by orientation contrasts over occipital areas within 100 ms after stimulus onset. These contrast effects occurred 10–20 ms later than the C1 components evoked by identically located abrupt onset stimuli which indexes the initial feedforward activity in V1. Compared with those in the UVF, orientation contrasts in the LVF evoked earlier and stronger activities, probably reflecting a LVF advantage in processing of orientation contrast. Even when orientation contrasts were rendered almost invisible by backward masking (in Experiment 2), the early contrast effect in the LVF was not disrupted. These findings imply that automatic processing of orientation contrast could occur at early visual cortical processing stages, but was slightly later than the initial feedforward processing in human V1; such automatic processing may involve either recurrent processing in V1 or feedforward processing in early extrastriate visual cortex.Highlights -We examined the earliest automatic processing of orientation contrast in humans with ERPs.-Significant orientation contrast effect started within 100 ms in early visual areas.-The earliest orientation contrast effect occurred later than the C1 evoked by abrupt onset stimuli.-The earliest orientation contrast effect was independent of top-down attention and awareness.-Automatic detection of orientation contrast arises slightly after the initial feedforward processing in V1.

Contextual Effects of High Dynamic Range (HDR) Luminance on Orientation Discrimination

Contextual cues are known to affect orientation discrimination and brightness perception, but how these two modalities interact in real-world perception is poorly understood. Studies of brightness perception based on laboratory stimuli are typically at ~100:1 luminance contrast ratio ('standard dynamic range', SDR), but real-world scenes have contrast ratios up to 1,000,000:1, and recent reports suggest that, at contrast ratios over 1000:1 ('high dynamic range', HDR), luminance normalization expands the perceived shadings of gray at the mode of the luminance distribution (Allred et al 2012). We hypothesized that the contextual mechanisms of luminance normalization and feature processing interact at the earliest stage of visual cortical processing, in primary visual cortex. We are measuring EEG, eye tracking, and visual recognition behavior under a two-alternative forced choice (2AFC) task and under rapid serial visual presentation (RSVP, 0.5-2 Hz). In both tasks, a contrast mixture of two orthogonal Gabors appears as a central target, and the subject is instructed to indicate via keypress the orientation with the stronger contrast. The target is surrounded by a 5 x 5 grid of Gabor flankers, each on a separate luminance patch from an HDR luminance distribution. In a baseline condition with uniform flanker orientation and isoluminant patches, we reliably reproduced the classic 'pop-out' effect (Li 1999). We predict that, at HDR luminance, the contextual orientation effect is stronger for patches that are more similar to the target luminance. We also predict that this luminance dependence is stronger for HDR than for SDR luminance distributions. We predict that the ERPs will be consistent with the integration of orientation and luminance cues in visual cortex and that the RSVP task will reveal the spatial dependency of this contextual effect. Meeting abstract presented at VSS 2018

Prior object-knowledge sharpens properties of early visual feature-detectors

Early stages of visual processing are carried out by neural circuits activated by simple and specific features, such as the orientation of an edge. A fundamental question in human vision is how the brain organises such intrinsically local information into meaningful properties of objects. Classic models of visual processing emphasise a one-directional flow of information from early feature-detectors to higher-level information-processing. By contrast to this view, and in line with predictive-coding models of perception, here, we provide evidence from human vision that high-level object representations dynamically interact with the earliest stages of cortical visual processing. In two experiments, we used ambiguous stimuli that, depending on the observer’s prior object-knowledge, can be perceived as either coherent objects or as a collection of meaningless patches. By manipulating object knowledge we were able to determine its impact on processing of low-level features while keeping sensory stimulation identical. Both studies demonstrate that perception of local features is facilitated in a manner consistent with an observer’s high-level object representation (i.e., with no effect on object-inconsistent features). Our results cannot be ascribed to attentional influences. Rather, they suggest that high-level object representations interact with and sharpen early feature-detectors, optimising their performance for the current perceptual context.

EEG Dynamics and Neural Generators in Implicit Navigational Image Processing in Adults with ADHD

In contrast to childhood ADHD that is characterized by inattention, impulsivity and hyperactivity, most adults with ADHD predominantly exhibit inattention. We used a new oddball paradigm using implicit navigational images and analyzed EEG dynamics with swLORETA inverse modeling of the evoked potential generators to study cortical processing in adults with ADHD and age-matched controls. In passive observation, we demonstrated that P350 amplitude, alpha–beta oscillation event-related synchronization (ERS) anticipation, and beta event-related desynchronization (ERD) were significantly smaller in ADHD. In the active condition, P100 duration was reduced and N140 amplitude increased for both deviant and frequent conditions in the ADHD. Alpha ERS and delta-theta ERS were reduced in the ADHD in the deviant condition. The left somatosensory area (BA2) and the right parietal lobe (BA31, BA40) contributed more to the P100 generators in the control than in the ADHD group, while the left frontal lobe (BA10) contributed more to the P100 generators in the ADHD. The left inferior parietal lobe (BA40) contributed more to the N140 generators in the control than the ADHD group while the right posterior cingulate (BA30) contributed more to the N140 generators in the ADHD. These findings reinforce the notion that earlier cortical stages of visual processing are compromised in adult ADHD by inducing the emergence of different even-related potential generators and EEG dynamics in ADHD. Considering that classical approaches for ADHD diagnosis are based on qualitative clinical investigation possibly biased by subjectivity, EEG analysis is another objective tool that might contribute to diagnosis, future neurofeedback or brain stimulation therapies.

Spatial Information in a Non-retinotopic Visual Cortex

Turtle dorsal cortex (dCx), a three-layered cortical area of the reptilian telencephalon, receives inputs from the retina via the thalamic lateral geniculate nucleus and constitutes the first cortical stage of visual processing. The receptive fields of dCx neurons usually occupy the entire contralateral visual field. Electrophysiological recordings in awake and anesthetized animals reveal that dCx is sensitive to the spatial structure of natural images, that dCx receptive fields are not entirely uniform across space, and that adaptation to repeated stimulation is position specific. Hence, spatial information can be found both at the single-neuron and population scales. Anatomical data are consistent with the absence of a clear retinotopic mapping of thalamocortical projections. The mapping and representation of visual space in this three-layered cortex thus differ from those found in mammalian primary visual cortex. Our results support the notion that dCx performs a global, rather than local, analysis of the visual scene.

Non-uniform phase sensitivity in spatial frequency maps of the human visual cortex.

Just as a portrait painting can come from a collection of coarse and fine details, natural vision can be decomposed into coarse and fine components. Previous studies have shown that the early visual areas in the brain represent these components in a map-like fashion. Other studies have shown that these same visual areas can be sensitive to how coarse and fine features line up in space. We found that the brain actually jointly represents both the scale of the feature (fine, medium, or coarse) and the alignment of these features in space. The results suggest that the visual cortex has an optimized representation particularly for the alignment of fine details, which are crucial in understanding the visual scene. Complex natural scenes can be decomposed into their oriented spatial frequency (SF) and phase relationships, both of which are represented locally at the earliest stages of cortical visual processing. The SF preference map in the human cortex, obtained using synthetic stimuli, is orderly and correlates strongly with eccentricity. In addition, early visual areas show sensitivity to the phase information that describes the relationship between SFs and thereby dictates the structure of the image. Taken together, two possibilities arise for the joint representation of SF and phase: either the entirety of the cortical SF map is uniformly sensitive to phase, or a particular set of SFs is selectively phase sensitive - for example, greater phase sensitivity for higher SFs that define fine-scale edges in a complex scene. To test between these two possibilities, we constructed a novel continuous natural scene video whereby phase information was maintained in one SF band but scrambled elsewhere. By shifting the central frequency of the phase-aligned band in time, we mapped the phase-sensitive SF preference of the visual cortex. Using functional magnetic resonance imaging, we found that phase sensitivity in early visual areas is biased toward higher SFs. Compared to a SF map of the same scene obtained using linear-filtered stimuli, a much larger patch of areas V1 and V2 is sensitive to the phase alignment of higher SFs. The results of early areas cannot be explained by attention. Our results suggest non-uniform sensitivity to phase alignment in population-level SF representations, with phase alignment being particularly important for fine-scale edge representations of natural scenes.

Temporal Information Entropy of the Blood-Oxygenation Level-Dependent Signals Increases in the Activated Human Primary Visual Cortex.

Time-domain analysis of blood-oxygenation level-dependent (BOLD) signals allows the identification of clusters of voxels responding to photic stimulation in primary visual cortex (V1). However, the characterization of information encoding into temporal properties of the BOLD signals of an activated cluster is poorly investigated. Here, we used Shannon entropy to determine spatial and temporal information encoding in the BOLD signal within the most strongly activated area of the human visual cortex during a hemifield photic stimulation. We determined the distribution profile of BOLD signals during epochs at rest and under stimulation within small (19–121 voxels) clusters designed to include only voxels driven by the stimulus as highly and uniformly as possible. We found consistent and significant increases (2–4% on average) in temporal information entropy during activation in contralateral but not ipsilateral V1, which was mirrored by an expected loss of spatial information entropy. These opposite changes coexisted with increases in both spatial and temporal mutual information (i.e., dependence) in contralateral V1. Thus, we showed that the first cortical stage of visual processing is characterized by a specific spatiotemporal rearrangement of intracluster BOLD responses. Our results indicate that while in the space domain BOLD maps may be incapable of capturing the functional specialization of small neuronal populations due to relatively low spatial resolution, some information encoding may still be revealed in the temporal domain by an increase of temporal information entropy.

A Comparison of Visual Response Properties in the Lateral Geniculate Nucleus and Primary Visual Cortex of Awake and Anesthetized Mice

The primary visual cortex (V1) of the mouse and its inputs from visual thalamus (dLGN), have become a dominant model for studying information processing in the brain. Early surveys of visual response properties (receptive fields) were performed in anesthetized animals. Although most recent studies of V1 have been performed in awake animals to examine links between vision and behavior, there have been few comprehensive studies of receptive field properties in the awake mouse, especially in dLGN and lateral posterior nucleus. We have performed a comparative survey of receptive fields in dLGN, lateral posterior nucleus, and V1 in anesthetized and awake mice. We found multiple differences in processing of time-varying stimuli, whereas the spatial aspects of receptive fields remain comparatively unchanged.

How a New Color Aftereffect Happened To Turn Up at Oberlin, 50 Years Ago

At Oberlin in the ‘60s, Introductory Psychology was a full-year course introducing general experimental psychology as a biosocial science. Single cell recordings from cat and monkey visual cortex had recently enabled visual science to move beyond the retina to the early cortical stages of visual processing. Ivo Kohler had reported studies of behavior during continuous wearing of prism spectacles that displace the retinal image laterally. They showed that one can regain fairly normal visuomotor coordination in a few weeks. His reports made adaptation to prism glasses a hot topic for investigation.

The role of early stages of cortical visual processing in size and distance judgment: A transcranial direct current stimulation study

Recent research suggests that V1 plays an active role in the judgment of size and distance. Nevertheless, no research has been performed using direct brain stimulation to address this issue. We used transcranial direct-current stimulation (tDCS) to directly modulate the early stages of cortical visual processing while measuring size and distance perception with a psychophysical scaling method of magnitude estimation in a repeated-measures design. The subjects randomly received anodal, cathodal, and sham tDCS in separate sessions starting with size or distance judgment tasks. Power functions were fit to the size judgment data, whereas logarithmic functions were fit to distance judgment data. Slopes and R2 were compared with separate repeated-measures analyses of variance with two factors: task (size vs. distance) and tDCS (anodal vs. cathodal vs. sham). Anodal tDCS significantly decreased slopes, apparently interfering with size perception. No effects were found for distance perception. Consistent with previous studies, the results of the size task appeared to reflect a prothetic continuum, whereas the results of the distance task seemed to reflect a metathetic continuum. The differential effects of tDCS on these tasks may support the hypothesis that different physiological mechanisms underlie judgments on these two continua. The results further suggest the complex involvement of the early visual cortex in size judgment tasks that go beyond the simple representation of low-level stimulus properties. This supports predictive coding models and experimental findings that suggest that higher-order visual areas may inhibit incoming information from the early visual cortex through feedback connections when complex tasks are performed.

Source Localisation of Visual Evoked Potentials in Congenitally Deaf Individuals

Previous studies have suggested that individuals deprived of auditory input can compensate with specific superior abilities in the remaining sensory modalities. To better understand the neural basis of deafness-induced changes, the present study used electroencephalography to examine visual functions and cross-modal reorganization of the auditory cortex in deaf individuals. Congenitally deaf participants and hearing controls were presented with reversing chequerboard stimuli that were systematically modulated in luminance ratio. The two groups of participants showed similar modulation of visual evoked potential (VEP) amplitudes (N85, P110) and latencies (P110) as a function of luminance ratio. Analysis of VEPs revealed faster neural processing in deaf participants compared with hearing controls at early stages of cortical visual processing (N85). Deaf participants also showed higher amplitudes (P110) than hearing participants. In contrast to our expectations, the results from VEP source analysis revealed no clear evidence for cross-modal reorganization in the auditory cortex of deaf participants. However, deaf participants tended to show higher activation in posterior parietal cortex (PPC). Moreover, modulation of PPC responses as a function of luminance was also stronger in deaf than in hearing participants. Taken together, these findings are an indication of more efficient neural processing of visual information in the deaf, which may relate to functional changes, in particular in multisensory parietal cortex, as a consequence of early auditory deprivation.

Spatial limitations of fast temporal segmentation are best modeled by V1 receptive fields

The fine temporal structure of events influences the spatial grouping and segmentation of visual-scene elements. Although adjacent regions flickering asynchronously at high temporal frequencies appear identical, the visual system signals a boundary between them. These "phantom contours" disappear when the gap between regions exceeds a critical value (g(max)). We used g(max) as an index of neuronal receptive-field size to compare with known receptive-field data from along the visual pathway and thus infer the location of the mechanism responsible for fast temporal segmentation. Observers viewed a circular stimulus reversing in luminance contrast at 20 Hz for 500 ms. A gap of constant retinal eccentricity segmented each stimulus quadrant; on each trial, participants identified a target quadrant containing counterphasing inner and outer segments. Through varying the gap width, g(max) was determined at a range of retinal eccentricities. We found that g(max) increased from 0.3° to 0.8° for eccentricities from 2° to 12°. These values correspond to receptive-field diameters of neurons in primary visual cortex that have been reported in single-cell and fMRI studies and are consistent with the spatial limitations of motion detection. In a further experiment, we found that modulation sensitivity depended critically on the length of the contour and could be predicted by a simple model of spatial summation in early cortical neurons. The results suggest that temporal segmentation is achieved by neurons at the earliest cortical stages of visual processing, most likely in primary visual cortex.

Earliest stages of visual cortical processing are not modified by attentional load.

This study investigated the effects of attentional load on neural responses to attended and irrelevant visual stimuli by recording high-density event-related potentials (ERPs) from the scalp in normal adult subjects. Peripheral (upper and lower visual field) and central stimuli were presented in random order at a rapid rate while subjects responded to targets among the central stimuli. Color detection and color-orientation conjunction search tasks were used as the low- and high-load tasks, respectively. Behavioral results showed significant load effects on both accuracy and reaction time for target detections. ERP results revealed no significant load effect on the initial C1 component (60-100 ms) evoked by either central-relevant or peripheral-irrelevant stimuli. Source analysis with dipole modeling confirmed previous reports that the C1 includes the initial evoked response in primary visual cortex. Source analyses indicated that high attentional load enhanced the early (70-140 ms) neural response to central-relevant stimuli in ventral-lateral extrastriate cortex, whereas load effects on peripheral-irrelevant stimulus processing started at 110 ms and were localized to more dorsal and anterior extrastriate cortical areas. These results provide evidence that the earliest stages of visual cortical processing are not modified by attentional load and show that attentional load affects the processing of task relevant and irrelevant stimuli in different ways.

Strabismus Disrupts Binocular Synaptic Integration in Primary Visual Cortex

Visual disruption early in development dramatically changes how primary visual cortex neurons integrate binocular inputs. The disruption is paradigmatic for investigating the synaptic basis of long-term changes in cortical function, because the primary visual cortex is the site of binocular convergence. The underlying alterations in circuitry by visual disruption remain poorly understood. Here we compare membrane potential responses, observed via whole-cell recordings in vivo, of primary visual cortex neurons in normal adult cats with those of cats in which strabismus was induced before the developmental critical period. In strabismic cats, we observed a dramatic shift in the ocular dominance distribution of simple cells, the first stage of visual cortical processing, toward responding to one eye instead of both, but not in complex cells, which receive inputs from simple cells. Both simple and complex cells no longer conveyed the binocular information needed for depth perception based on binocular cues. There was concomitant binocular suppression such that responses were weaker with binocular than with monocular stimulation. Our estimates of the excitatory and inhibitory input to single neurons indicate binocular suppression that was not evident in synaptic excitation, but arose de novo because of synaptic inhibition. Further constraints on circuit models of plasticity result from indications that the ratio of excitation to inhibition evoked by monocular stimulation decreased mainly for nonpreferred eye stimulation. Although we documented changes in synaptic input throughout primary visual cortex, a circuit model with plasticity at only thalamocortical synapses is sufficient to account for our observations.

Altered Contextual Modulation of Primary Visual Cortex Responses in Schizophrenia

Schizophrenia is typically associated with higher-level cognitive symptoms, such as disorganized thoughts, delusions, and hallucinations. However, deficits in visual processing have been consistently reported with the illness. Here, we provide strong neurophysiological evidence for a marked perturbation at the earliest level of cortical visual processing in patients with paranoid schizophrenia. Using functional magnetic resonance imaging (fMRI) and adapting a well-established approach from electrophysiology, we found that orientation-specific contextual modulation of cortical responses in human primary visual cortex (V1)--a hallmark of early neural encoding of visual stimuli--is dramatically reduced in patients with schizophrenia. This indicates that contextual processing in schizophrenia is altered at the earliest stages of visual cortical processing and supports current theories that emphasize the role of abnormalities in perceptual synthesis (eg, false inference) in schizophrenia.

How the Visual Cortex Handles Stimulus Noise: Insights from Amblyopia

Adding noise to a visual image makes object recognition more effortful and has a widespread effect on human electrophysiological responses. However, visual cortical processes directly involved in handling the stimulus noise have yet to be identified and dissociated from the modulation of the neural responses due to the deteriorated structural information and increased stimulus uncertainty in the case of noisy images. Here we show that the impairment of face gender categorization performance in the case of noisy images in amblyopic patients correlates with amblyopic deficits measured in the noise-induced modulation of the P1/P2 components of single-trial event-related potentials (ERP). On the other hand, the N170 ERP component is similarly affected by the presence of noise in the two eyes and its modulation does not predict the behavioral deficit. These results have revealed that the efficient processing of noisy images depends on the engagement of additional processing resources both at the early, feature-specific as well as later, object-level stages of visual cortical processing reflected in the P1 and P2 ERP components, respectively. Our findings also suggest that noise-induced modulation of the N170 component might reflect diminished face-selective neuronal responses to face images with deteriorated structural information.