Customizing spatial remapping of letters to aid reading in the presence of a simulated central field loss.

Reading is a primary concern of patients with central field loss (CFL) because it is typically performed with foveal vision. Spatial remapping offers one potential avenue to aid in reading; it entails shifting occluded letters to retinal areas where vision is functional. Here, we introduce a method of creating and testing different remapping strategies-ways to remap text-customized for CFL of different shapes. By simulating CFL in typically-sighted individuals, we tested the customization hypothesis-that the benefits of different remapping strategies will depend on the properties of the CFL. That is, remapping strategies will aid reading differentially in the presence of differently shaped CFL. In Experiment 1, letter recognition in the presence of differently shaped CFL was assessed in and around central vision. Using these letter recognition "maps" different spatial remappings were created and tested in Experiment 2 using a word recognition task. Results showed that the horizontal gap remapping, which did not remap any letters vertically, resulted in the best word recognition. Results were also consistent with the customization hypothesis; the benefits of different remappings on word recognition depended on the different CFL shapes. Although the horizontal gap remapping resulted in very good word recognition, tailoring remapping strategies to the shape of patients' CFL may aid reading with the wide range of sizes and shapes encountered by patients with CFL.

Adapting to Visual Noise Alleviates Visual Snow.

Purpose: Visual snow syndrome-characterized by flickering specks throughout the visual field and accompanied by other symptoms-can disrupt daily life and affects roughly 2% of the population. However, its neural bases remain mysterious, and treatments are lacking. Here, we report the first intervention that can temporarily eliminate the visual snow symptom, allowing many observers to see the world without snow for the first time since symptom onset. Prolonged viewing of a visual stimulus strongly reduces the responsiveness of the visual pathways to subsequent stimuli, and we tested whether such adaptation could affect visual snow. Methods: Participants with visual snow (total n = 27) viewed high-contrast dynamic noise patterns, resembling television static, and then judged the strength of the symptom. Results: Visual snow was temporarily reduced in strength to the point that it was invisible at longer adaptation durations for most observers. The effect followed typical trends of adaptation for physical stimuli in normally sighted observers: Effect duration increased monotonically with duration of exposure to the adapter and was specific to dynamic noise. Conclusions: These results establish that spontaneous neural activity in the visual system is causally related to the visual snow percept. Because they perceive this activity, people with visual snow may provide a unique window into the generation and suppression of noise in the visual system. Adaptation allows reliable experimental control over visual snow, and so is a strong candidate for diagnostic testing and a promising tool for further understanding its neural origins, which could in turn aid the development of treatments.

An Accumulating Neural Signal Underlying Binocular Rivalry Dynamics.

During binocular rivalry, conflicting images are presented one to each eye and perception alternates stochastically between them. Despite stable percepts between alternations, modeling suggests that neural signals representing the two images change gradually, and that the duration of stable percepts are determined by the time required for these signals to reach a threshold that triggers an alternation. However, direct physiological evidence for such signals has been lacking. Here, we identify a neural signal in the human visual cortex that shows these predicted properties. We measured steady-state visual evoked potentials (SSVEPs) in 84 human participants (62 females, 22 males) who were presented with orthogonal gratings, one to each eye, flickering at different frequencies. Participants indicated their percept while EEG data were collected. The time courses of the SSVEP amplitudes at the two frequencies were then compared across different percept durations, within participants. For all durations, the amplitude of signals corresponding to the suppressed stimulus increased and the amplitude corresponding to the dominant stimulus decreased throughout the percept. Critically, longer percepts were characterized by more gradual increases in the suppressed signal and more gradual decreases of the dominant signal. Changes in signals were similar and rapid at the end of all percepts, presumably reflecting perceptual transitions. These features of the SSVEP time courses are well predicted by a model in which perceptual transitions are produced by the accumulation of noisy signals. Identification of this signal underlying binocular rivalry should allow strong tests of neural models of rivalry, bistable perception, and neural suppression.SIGNIFICANCE STATEMENT During binocular rivalry, two conflicting images are presented to the two eyes and perception alternates between them, with switches occurring at seemingly random times. Rivalry is an important and longstanding model system in neuroscience, used for understanding neural suppression, intrinsic neural dynamics, and even the neural correlates of consciousness. All models of rivalry propose that it depends on gradually changing neural activity that on reaching some threshold triggers the perceptual switches. This manuscript reports the first physiological measurement of neural signals with that set of properties in human participants. The signals, measured with EEG in human observers, closely match the predictions of recent models of rivalry, and should pave the way for much future work.

Slowed alpha oscillations and percept formation in psychotic psychopathology.

Introduction: Psychosis is in part defined by disturbances in perception. Recent investigations have implicated the speed of alpha oscillations observed in brain electrical activity as reflective of a sampling rate of the visual environment and perception. Although both slowed alpha oscillations and aberrant percept formation are evident in disorders of psychotic psychopathology such as schizophrenia it is unclear whether slow alpha accounts for abnormal visual perception in these disorders. Methods: To examine the role of the speed of alpha oscillations in perception in psychotic psychopathology we gathered resting-state magneto-encephalography data from probands with psychotic psychopathology (i.e., schizophrenia, schizoaffective disorder, and bipolar disorder with a history of psychosis), their biological siblings, and healthy controls. We appraised visual perceptual function without the confound of cognitive ability and effort through the use of a simple binocular rivalry task. Results: We found a slowed pace of alpha oscillations in psychotic psychopathology that was associated with longer percept durations during binocular rivalry, consistent with the assertion that occipital alpha oscillations govern the rate of accumulation of visual information used to generate percepts. Alpha speed varied widely across individuals with psychotic psychopathology and was highly stable across several months indicating that it is likely a trait characteristic of neural function that is relevant to visual perception. Finally, a lower speed of alpha oscillation was associated with a lower IQ and greater disorder symptomatology implying that the effects of the endogenous neural oscillation on visual perception may have wider consequences for everyday functioning. Discussion: Slowed alpha oscillations in individuals with psychotic psychopathology appear to reflect altered neural functions related to percept formation.

Variability of the Surface Area of the V1, V2, and V3 Maps in a Large Sample of Human Observers.

How variable is the functionally defined structure of early visual areas in human cortex and how much variability is shared between twins? Here we quantify individual differences in the best understood functionally defined regions of cortex: V1, V2, V3. The Human Connectome Project 7T Retinotopy Dataset includes retinotopic measurements from 181 subjects (109 female, 72 male), including many twins. We trained four "anatomists" to manually define V1-V3 using retinotopic features. These definitions were more accurate than automated anatomical templates and showed that surface areas for these maps varied more than threefold across individuals. This threefold variation was little changed when normalizing visual area size by the surface area of the entire cerebral cortex. In addition to varying in size, we find that visual areas vary in how they sample the visual field. Specifically, the cortical magnification function differed substantially among individuals, with the relative amount of cortex devoted to central vision varying by more than a factor of 2. To complement the variability analysis, we examined the similarity of visual area size and structure across twins. Whereas the twin sample sizes are too small to make precise heritability estimates (50 monozygotic pairs, 34 dizygotic pairs), they nonetheless reveal high correlations, consistent with strong effects of the combination of shared genes and environment on visual area size. Collectively, these results provide the most comprehensive account of individual variability in visual area structure to date, and provide a robust population benchmark against which new individuals and developmental and clinical populations can be compared.SIGNIFICANCE STATEMENT Areas V1, V2, and V3 are among the best studied functionally defined regions in human cortex. Using the largest retinotopy dataset to date, we characterized the variability of these regions across individuals and the similarity between twin pairs. We find that the size of visual areas varies dramatically (up to 3.5×) across healthy young adults, far more than the variability of the cerebral cortex size as a whole. Much of this variability appears to arise from inherited factors, as we find very high correlations in visual area size between monozygotic twin pairs, and lower but still substantial correlations between dizygotic twin pairs. These results provide the most comprehensive assessment of how functionally defined visual cortex varies across the population to date.

Visual mode switching: Improved general compensation for environmental color changes requires only one exposure per day.

When the visual environment changes, vision adapts in order to maintain accurate perception. For repeatedly encountered environmental changes, the visual system may learn to adjust immediately, a process called "visual mode switching." For example, following experience with red glasses, participants report that the glasses' redness fades instantly when they put the glasses on. Here we tested (1) whether once-daily experience suffices for learning to switch visual modes and (2) whether effects of mode switching apply to most stimuli affected by the environmental change. In Experiment 1, 12 participants wore bright red glasses for a single 5-hr period each day for 5 days, and we tested for changes in the perception of unique yellow, which contains neither red nor green. In Experiment 2, we tested how mode switching affects larger parts of the color space. Thirteen participants donned and removed the glasses multiple times a day for 5 days, and we used a dissimilarity rating task to measure and track perception of many different colors. Across days, immediately upon donning the glasses, the world appeared less and less reddish (Experiment 1), and colors across the whole color space appeared more and more normal (Experiment 2). These results indicate that mode switching can be acquired from a once-daily experience, and it applies to most stimuli in a given environment. These findings may help to predict when and how mode switching occurs outside the laboratory.

Visual adaptation selective for individual limbs reveals hierarchical human body representation.

The spatial relationships between body parts are a rich source of information for person perception, with even simple pairs of parts providing highly valuable information. Computation of these relationships would benefit from a hierarchical representation, where body parts are represented individually. We hypothesized that the human visual system makes use of such representations. To test this hypothesis, we used adaptation to determine whether observers were sensitive to changes in the length of one body part relative to another. Observers viewed forearm/upper arm pairs where the forearm had been either lengthened or shortened, judging the perceived length of the forearm. Observers then adapted to a variety of different stimuli (e.g., arms, objects, etc.) in different orientations and visual field locations. We found that following adaptation to distorted limbs, but not non-limb objects, observers experienced a shift in perceived forearm length. Furthermore, this effect partially transferred across different orientations and visual field locations. Taken together, these results suggest the effect arises in high level mechanisms specialized for specific body parts, providing evidence for a representation of bodies based on parts and their relationships.

Benefits of deep learning classification of continuous noninvasive brain-computer interface control.

Objective. Noninvasive brain-computer interfaces (BCIs) assist paralyzed patients by providing access to the world without requiring surgical intervention. Prior work has suggested that EEG motor imagery based BCI can benefit from increased decoding accuracy through the application of deep learning methods, such as convolutional neural networks (CNNs).Approach. Here, we examine whether these improvements can generalize to practical scenarios such as continuous control tasks (as opposed to prior work reporting one classification per trial), whether valuable information remains latent outside of the motor cortex (as no prior work has compared full scalp coverage to motor only electrode montages), and the existing challenges to the practical implementation of deep-learning based continuous BCI control.Main results. We report that: (1) deep learning methods significantly increase offline performance compared to standard methods on an independent, large, and longitudinal online motor imagery BCI dataset with up to 4-classes and continuous 2D feedback; (2) our results suggest that a variety of neural biomarkers for BCI, including those outside the motor cortex, can be detected and used to improve performance through deep learning methods, and (3) tuning neural network output will be an important step in optimizing online BCI control, as we found the CNN models trained with full scalp EEG also significantly reduce the average trial length in a simulated online cursor control environment.Significance. This work demonstrates the benefits of CNNs classification during BCI control while providing evidence that electrode montage selection and the mapping of CNN output to device control will be important design choices in CNN based BCIs.

Continuous sensorimotor rhythm based brain computer interface learning in a large population.

Brain computer interfaces (BCIs) are valuable tools that expand the nature of communication through bypassing traditional neuromuscular pathways. The non-invasive, intuitive, and continuous nature of sensorimotor rhythm (SMR) based BCIs enables individuals to control computers, robotic arms, wheel-chairs, and even drones by decoding motor imagination from electroencephalography (EEG). Large and uniform datasets are needed to design, evaluate, and improve the BCI algorithms. In this work, we release a large and longitudinal dataset collected during a study that examined how individuals learn to control SMR-BCIs. The dataset contains over 600 hours of EEG recordings collected during online and continuous BCI control from 62 healthy adults, (mostly) right hand dominant participants, across (up to) 11 training sessions per participant. The data record consists of 598 recording sessions, and over 250,000 trials of 4 different motor-imagery-based BCI tasks. The current dataset presents one of the largest and most complex SMR-BCI datasets publicly available to date and should be useful for the development of improved algorithms for BCI control.

The McCollough World: Induction of orientation-contingent aftereffects with an altered-reality system.

The McCollough Effect is a color aftereffect produced by exposure to colored, oriented patterns. For example, following adaptation to vertical red and horizontal green stripes in alternation, vertical black and white patterns appear greenish, while horizontal black and white patterns appear reddish. The striking aspect of the McCollough Effect is that just a few minutes of adaptation can produce an aftereffect lasting days or weeks. Though this effect is easily induced, previous work has shown that stronger effects can be achieved with longer periods of adaptation. To allow especially long adaptation durations, the current work develops a novel method of induction of the McCollough Effect using live video feed, filtered by orientation, and viewed with a head-mounted display. Results showed that this "McCollough World" paradigm was as strong an inducer (per unit time) as traditional paradigms using gratings, while allowing observers to adapt comfortably for multiple hours. Two hours of McCollough World adaptation produced effects that were significantly larger than 20 min of traditional adaptation, which is close to the tolerance limits for gratings. This work provides insight into the features necessary for induction of the McCollough Effect and provides a strategy for creating especially strong and long-lasting color aftereffects.

Color Compensation in Anomalous Trichromats Assessed with fMRI.

Anomalous trichromacy is a common form of congenital color deficiency resulting from a genetic alteration in the photopigments of the eye's light receptors. The changes reduce sensitivity to reddish and greenish hues, yet previous work suggests that these observers may experience the world to be more colorful than their altered receptor sensitivities would predict, potentially indicating an amplification of post-receptoral signals. However, past evidence suggesting such a gain adjustment rests on subjective measures of color appearance or salience. We directly tested for neural amplification by using fMRI to measure cortical responses in color-anomalous and normal control observers. Color contrast response functions were measured in two experiments with different tasks to control for attentional factors. Both experiments showed a predictable reduction in chromatic responses for anomalous trichromats in primary visual cortex. However, in later areas V2v and V3v, chromatic responses in the two groups were indistinguishable. Our results provide direct evidence for neural plasticity that compensates for the deficiency in the initial receptor color signals and suggest that the site of this compensation is in early visual cortex.

Depth-dependent functional MRI responses to chromatic and achromatic stimuli throughout V1 and V2.

In the primate visual system, form (shape, location) and color information are processed in separate but interacting pathways. Recent access to high-resolution neuroimaging has facilitated the exploration of the structure of these pathways at the mesoscopic level in the human visual cortex. We used 7T fMRI to observe selective activation of the primary visual cortex to chromatic versus achromatic stimuli in five participants across two scanning sessions. Achromatic checkerboards with low spatial frequency and high temporal frequency targeted the color-insensitive magnocellular pathway. Chromatic checkerboards with higher spatial frequency and low temporal frequency targeted the color-selective parvocellular pathway. This work resulted in three main findings. First, responses driven by chromatic stimuli had a laminar profile biased towards superficial layers of V1, as compared to responses driven by achromatic stimuli. Second, we found stronger preference for chromatic stimuli in parafoveal V1 compared with peripheral V1. Finally, we found alternating, stimulus-selective bands stemming from the V1 border into V2 and V3. Similar alternating patterns have been previously found in both NHP and human extrastriate cortex. Together, our findings confirm the utility of fMRI for revealing details of mesoscopic neural architecture in human cortex.

Visual mode switching learned through repeated adaptation to color.

When the environment changes, vision adapts to maintain accurate perception. For repeatedly encountered environments, learning to adjust more rapidly would be beneficial, but past work remains inconclusive. We tested if the visual system can learn such visual mode switching for a strongly color-tinted environment, where adaptation causes the dominant hue to fade over time. Eleven observers wore bright red glasses for five 1-hr periods per day, for 5 days. Color adaptation was measured by asking observers to identify 'unique yellow', appearing neither reddish nor greenish. As expected, the world appeared less and less reddish during the 1-hr periods of glasses wear. Critically, across days the world also appeared significantly less reddish immediately upon donning the glasses. These results indicate that the visual system learned to rapidly adjust to the reddish environment, switching modes to stabilize color vision. Mode switching likely provides a general strategy to optimize perceptual processes.

Structural and Functional Connectivity of Visual Cortex in Schizophrenia and Bipolar Disorder: A Graph-Theoretic Analysis.

Visual processing abnormalities in schizophrenia (SZ) are poorly understood, yet predict functional outcomes in the disorder. Bipolar disorder (BD) may involve similar visual processing deficits. Converging evidence suggests that visual processing may be relatively normal at early stages of visual processing such as early visual cortex (EVC), but that processing abnormalities may become more pronounced by mid-level visual areas such as lateral occipital cortex (LO). However, little is known about the connectivity of the visual system in SZ and BD. If the flow of information to, from, or within the visual system is disrupted by reduced connectivity, this could help to explain perceptual deficits. In the present study, we performed a targeted analysis of the structural and functional connectivity of the visual system using graph-theoretic metrics in a sample of 48 SZ, 46 BD, and 47 control participants. Specifically, we calculated parallel measures of local efficiency for EVC and LO from both diffusion weighted imaging data (structural) and resting-state (functional) imaging data. We found no structural connectivity differences between the groups. However, there was a significant group difference in functional connectivity and a significant group-by-region interaction driven by reduced LO connectivity in SZ relative to HC, whereas BD was approximately intermediate to the other 2 groups. We replicated this pattern of results using a different brain atlas. These findings support and extend theoretical models of perceptual dysfunction in SZ, providing a framework for further investigation of visual deficits linked to functional outcomes in SZ and related disorders.

Higher-Level Meta-Adaptation Mitigates Visual Distortions Produced by Lower-Level Adaptation.

The visual system adapts to the environment, changing neural responses to aid efficiency and improve perception. However, these changes sometimes lead to negative consequences: If neurons at later processing stages fail to account for adaptation at earlier stages, perceptual errors result, including common visual illusions. These negative effects of adaptation have been termed the coding catastrophe. How does the visual system resolve them? We hypothesized that higher-level adaptation can correct errors arising from the coding catastrophe by changing what appears normal, a common form of adaptation across domains. Observers (N = 15) viewed flickering checkerboards that caused a normal face to appear distorted. We tested whether the visual system can adapt to this adaptation-distorted face through repeated viewing. Results from two experiments show that such meta-adaptation does occur and that it makes the distorted face gradually appear more normal. Meta-adaptation may be a general strategy to correct negative consequences of low-level adaptation.

Natural-scene-based Steady-state Visual Evoked Potentials Reveal Effects of Short-term Monocular Deprivation.

Ocular dominance plasticity beyond the critical period has been demonstrated in adult humans in recent investigations of short-term monocular deprivation (MD). To our knowledge, all previous research adopted non-natural synthetic stimuli in testing perceptual ocular dominance before and after the MD. However, it is recognized that complex natural stimuli may engage cortical mechanisms substantially different from simple synthetic stimuli. Therefore, it remains largely unknown whether reshaping of ocular dominance following MD could be observed during perception of natural scene stimuli without conspicuous interocular competition. Here we used the steady-state visual evoked potential (SSVEP) technique to measure the ocular-specific neural effects of MD with natural scene stimuli where the two eyes' images were tagged with different frequencies. Two hours of MD boosted the neural gain for the deprived eye. During the course of MD, the SSVEP amplitude ratio for the deprived eye compared to the non-deprived eye increased significantly over time, indicating a progressive increase of neural gain for the deprived eye. These findings demonstrate that the effects of short-term MD can manifest when viewing natural scenes, providing a natural case in support of the homeostatic compensation theory of MD. Our work also indicates that the technique of natural-scene-based SSVEP could be particularly useful for future work exploring the neural dynamics during adaptation to natural stimuli.

Control of visual adaptation depends upon task.

The visual system optimizes its functioning for a given environment through processes collectively called adaptation. It is currently unknown, however, whether adaptation is affected by the particular task the observer performs within that environment. Two experiments tested whether this is the case. Observers adapted to high contrast grating patterns, and the decay of adaptation was measured using a version of the tilt-aftereffect, while they performed two different secondary tasks. One task involved judging the luminance of a small circular spot at fixation, and was expected to be unaffected by adaptation. The other secondary task involved judging a low contrast grating, and adaptation was expected to make this task difficult by reducing the visibility of the grating. Identical displays containing both a fixation spot and a grating were used for both tasks. Tilt-aftereffects were smaller when subjects concurrently performed the grating task than when they performed the fixation task. These results suggest that the control of adaptation, in this case its decay, is sensitive to the nature of the task the observer is performing. Adaptation may attempt to optimize vision with respect to many different criteria simultaneously; task is likely one of the criteria included in this process.

Evidence for intact stimulus-specific neural adaptation for visual objects in schizophrenia and bipolar disorder: An ERP study.

People with schizophrenia (SZ) or bipolar disorder (BD) experience dysfunction in visual processing. Dysfunctional neural tuning, in which neurons and neuronal populations are selectively activated by specific features of visual stimuli, may contribute to these deficits. Few studies have examined this possibility and there are inconsistent findings of tuning deficits in the literature. We utilized an event-related potential (ERP) paradigm to examine neural adaptation for visual objects, a measure of neural tuning whereby neurons respond less strongly to the repeated presentation of the same stimulus. Seventy-seven SZ, 53 BD, and 49 healthy comparison participants (HC) were examined. In three separate conditions, pictures of objects were presented repeatedly: the same object (SS), different objects from the same category (e.g., two different vases; SD), or different objects from different categories (e.g., a barrel and a clock, DD). Mass-univariate cluster-based permutation analyses identified electrodes and time-windows in which there were significant differences between the SS vs. DD and the SD vs. DD conditions. Mean ERP amplitudes were extracted from these clusters and analyzed for group differences. Results revealed a significant condition difference over parieto-occipital electrodes for the SS-DD comparison between 109-164 ms and for the SD-DD comparison between 78-203 ms, with larger amplitudes in the DD compared to either SS or SD condition. However, there were no significant differences in the pattern of results between groups. Thus, while we found neural adaptation effects using this ERP paradigm, we did not find evidence of group differences. Our results suggest that people with SZ or BD may not exhibit deficits in neural tuning for processing of visual objects using this EEG task with rapidly presented stimuli. However, the results are inconsistent with other studies using different methodologies (e.g., fMRI, behavioral tasks) that have found tuning deficits in people with schizophrenia.

Frequency of alpha oscillation predicts individual differences in perceptual stability during binocular rivalry.

When ambiguous visual stimuli have multiple interpretations, human perception can alternate between them, producing perceptual multistability. There is a large variation between individuals in how long stable percepts endure, on average, between switches, but the underlying neural basis of this individual difference in perceptual dynamics remains obscure. Here, we show that in one widely studied multistable paradigm-binocular rivalry-perceptual stability in individuals is predicted by the frequency of their neural oscillations within the alpha range (7-13 Hz). Our results suggest revising models of rivalry to incorporate effects of neural oscillations on perceptual alternations, and raise the possibility that a common factor may influence dynamics in many neural processes.

fMRI evidence of aberrant neural adaptation for objects in schizophrenia and bipolar disorder.

Functional magnetic resonance imaging (fMRI) adaptation (also known as fMRI repetition suppression) has been widely used to characterize stimulus selectivity in vivo, a fundamental feature of neuronal processing in the brain. We investigated whether SZ patients and BD patients show aberrant fMRI adaptation for object perception. About 52 SZ patients, 55 BD patients, and 53 community controls completed an object discrimination task with three conditions: the same object presented twice, two exemplars from the same category, and two exemplars from different categories. We also administered two functional localizer tasks. A region of interest analysis was employed to evaluate a priori hypotheses about the lateral occipital complex (LOC) and early visual cortex (EVC). An exploratory whole brain analysis was also conducted. In the LOC and EVC, controls showed the expected reduced fMRI responses to repeated presentation of the same objects compared with different objects (i.e., fMRI adaptation for objects, p < .001). SZ patients showed an adaptation effect that was significantly smaller compared with controls. BD patients showed a lack of fMRI adaptation. The whole brain analyses showed enhanced fMRI responses to repeated presentation of the same objects only in BD patients in several brain regions including anterior cingulate cortex. This study was the first to employ fMRI adaptation for objects in SZ and BD. The current findings provide empirical evidence of aberrant fMRI adaptation in the visual cortex in SZ and BD, but in distinctly different ways.