Breaking the circularity in circular analyses: Simulations and formal treatment of the flattened average approach.

There has been considerable debate and concern as to whether there is a replication crisis in the scientific literature. A likely cause of poor replication is the multiple comparisons problem. An important way in which this problem can manifest in the M/EEG context is through post hoc tailoring of analysis windows (a.k.a. regions-of-interest, ROIs) to landmarks in the collected data. Post hoc tailoring of ROIs is used because it allows researchers to adapt to inter-experiment variability and discover novel differences that fall outside of windows defined by prior precedent, thereby reducing Type II errors. However, this approach can dramatically inflate Type I error rates. One way to avoid this problem is to tailor windows according to a contrast that is orthogonal (strictly parametrically orthogonal) to the contrast being tested. A key approach of this kind is to identify windows on a fully flattened average. On the basis of simulations, this approach has been argued to be safe for post hoc tailoring of analysis windows under many conditions. Here, we present further simulations and mathematical proofs to show exactly why the Fully Flattened Average approach is unbiased, providing a formal grounding to the approach, clarifying the limits of its applicability and resolving published misconceptions about the method. We also provide a statistical power analysis, which shows that, in specific contexts, the fully flattened average approach provides higher statistical power than Fieldtrip cluster inference. This suggests that the Fully Flattened Average approach will enable researchers to identify more effects from their data without incurring an inflation of the false positive rate.

Response dependence of reversal-related ERP components in perception of ambiguous figures.

Perceptual multi-stability is characterized by alternating interpretations of an unchanging stimulus input. The reversal negativity (RN) and reversal positivity (RP) ERP components show differences in electrophysiological responses between trials on which participants experience a perceptual reversal of a multi-stable stimulus versus trials without a reversal (i.e., stable). However, it is unclear to what extent these two ERP components reflect reversal-related perceptual processing rather than task and response processes. To address this, we varied task and response requirements while measuring the RN and RP. In the standard reversal task, participants indicated whether they saw a perceptual reversal on each trial. In contrast, in the identity task participants reported perceived identity of the stimulus (e.g., face or vase) without any reference to reversals. In some blocks, reversal trials required a response whereas in other blocks stable trials required a response. We found that the RN appeared independently of task and response style. However, the early latency RP component was only present when participants responded manually. For non-response trials, a component was found during the same latency as the RP but with inverted polarity. Our results suggest that the early RP component is dependent on response-related processes rather than being a pure neural signature of perceptual processes related to endogenous perceptual reversals.

Theta Phase-dependent Modulation of Perception by Concurrent Transcranial Alternating Current Stimulation and Periodic Visual Stimulation.

Sensory perception can be modulated by the phase of neural oscillations, especially in the theta and alpha ranges. Oscillatory activity in the visual cortex can be entrained by transcranial alternating current stimulation (tACS) as well as periodic visual stimulation (i.e., flicker). Combined tACS and visual flicker stimulation modulates BOLD response, and concurrent 4-Hz auditory click train, and tACS modulate auditory perception in a phase-dependent way. In this study, we investigated whether phase synchrony between concurrent tACS and periodic visual stimulation (i.e., flicker) can modulate performance on a visual matching task. Participants completed a visual matching task on a flickering visual stimulus while receiving either in-phase (0°) or asynchronous (180°, 90°, or 270°) tACS at alpha or theta frequency. Stimulation was applied over either occipital cortex or dorsolateral pFC. Visual performance was significantly better during theta frequency tACS over the visual cortex when it was in-phase (0°) with visual stimulus flicker, compared with antiphase (180°). This effect did not appear with alpha frequency flicker or with dorsolateral pFC stimulation. Furthermore, a control sham group showed no effect. There were no significant performance differences among the asynchronous (180°, 90°, and 270°) phase conditions. Extending previous studies on visual and auditory perception, our results support a crucial role of oscillatory phase in sensory perception and demonstrate a behaviorally relevant combination of visual flicker and tACS. The spatial and frequency specificity of our results have implications for research on the functional organization of perception.

Data-driven region-of-interest selection without inflating Type I error rate.

In ERP and other large multidimensional neuroscience data sets, researchers often select regions of interest (ROIs) for analysis. The method of ROI selection can critically affect the conclusions of a study by causing the researcher to miss effects in the data or to detect spurious effects. In practice, to avoid inflating Type I error rate (i.e., false positives), ROIs are often based on a priori hypotheses or independent information. However, this can be insensitive to experiment-specific variations in effect location (e.g., latency shifts) reducing power to detect effects. Data-driven ROI selection, in contrast, is nonindependent and uses the data under analysis to determine ROI positions. Therefore, it has potential to select ROIs based on experiment-specific information and increase power for detecting effects. However, data-driven methods have been criticized because they can substantially inflate Type I error rate. Here, we demonstrate, using simulations of simple ERP experiments, that data-driven ROI selection can indeed be more powerful than a priori hypotheses or independent information. Furthermore, we show that data-driven ROI selection using the aggregate grand average from trials (AGAT), despite being based on the data at hand, can be safely used for ROI selection under many circumstances. However, when there is a noise difference between conditions, using the AGAT can inflate Type I error and should be avoided. We identify critical assumptions for use of the AGAT and provide a basis for researchers to use, and reviewers to assess, data-driven methods of ROI localization in ERP and other studies.

Evidence for view-invariant face recognition units in unfamiliar face learning.

Many models of face recognition incorporate the idea of a face recognition unit (FRU), an abstracted representation formed from each experience of a face which aids recognition under novel viewing conditions. Some previous studies have failed to find evidence of this FRU representation. Here, we report three experiments which investigated this theoretical construct by modifying the face learning procedure from that in previous work. During learning, one or two views of previously unfamiliar faces were shown to participants in a serial matching task. Later, participants attempted to recognize both seen and novel views of the learned faces (recognition phase). Experiment 1 tested participants' recognition of a novel view, a day after learning. Experiment 2 was identical, but tested participants on the same day as learning. Experiment 3 repeated Experiment 1, but tested participants on a novel view that was outside the rotation of those views learned. Results revealed a significant advantage, across all experiments, for recognizing a novel view when two views had been learned compared to single view learning. The observed view invariance supports the notion that an FRU representation is established during multi-view face learning under particular learning conditions.

Causal evidence that intrinsic beta-frequency is relevant for enhanced signal propagation in the motor system as shown through rhythmic TMS.

Correlative evidence provides support for the idea that brain oscillations underpin neural computations. Recent work using rhythmic stimulation techniques in humans provide causal evidence but the interactions of these external signals with intrinsic rhythmicity remain unclear. Here, we show that sensorimotor cortex follows externally applied rhythmic TMS (rTMS) stimulation in the beta-band but that the elicited responses are strongest at the intrinsic individual beta peak frequency. While these entrainment effects are of short duration, even subthreshold rTMS pulses propagate through the network and elicit significant cortico-spinal coupling, particularly when stimulated at the individual beta-frequency. Our results show that externally enforced rhythmicity interacts with intrinsic brain rhythms such that the individual peak frequency determines the effect of rTMS. The observed downstream spinal effect at the resonance frequency provides evidence for the causal role of brain rhythms for signal propagation.

Training-induced recovery of low-level vision followed by mid-level perceptual improvements in developmental object and face agnosia.

Long-term deprivation of normal visual inputs can cause perceptual impairments at various levels of visual function, from basic visual acuity deficits, through mid-level deficits such as contour integration and motion coherence, to high-level face and object agnosia. Yet it is unclear whether training during adulthood, at a post-developmental stage of the adult visual system, can overcome such developmental impairments. Here, we visually trained LG, a developmental object and face agnosic individual. Prior to training, at the age of 20, LG's basic and mid-level visual functions such as visual acuity, crowding effects, and contour integration were underdeveloped relative to normal adult vision, corresponding to or poorer than those of 5-6 year olds (Gilaie-Dotan, Perry, Bonneh, Malach & Bentin, 2009). Intensive visual training, based on lateral interactions, was applied for a period of 9 months. LG's directly trained but also untrained visual functions such as visual acuity, crowding, binocular stereopsis and also mid-level contour integration improved significantly and reached near-age-level performance, with long-term (over 4 years) persistence. Moreover, mid-level functions that were tested post-training were found to be normal in LG. Some possible subtle improvement was observed in LG's higher-order visual functions such as object recognition and part integration, while LG's face perception skills have not improved thus far. These results suggest that corrective training at a post-developmental stage, even in the adult visual system, can prove effective, and its enduring effects are the basis for a revival of a developmental cascade that can lead to reduced perceptual impairments.

Preserved local but disrupted contextual figure-ground influences in an individual with abnormal function of intermediate visual areas.

Visual perception depends not only on local stimulus features but also on their relationship to the surrounding stimulus context, as evident in both local and contextual influences on figure-ground segmentation. Intermediate visual areas may play a role in such contextual influences, as we tested here by examining LG, a rare case of developmental visual agnosia. LG has no evident abnormality of brain structure and functional neuroimaging showed relatively normal V1 function, but his intermediate visual areas (V2/V3) function abnormally. We found that contextual influences on figure-ground organization were selectively disrupted in LG, while local sources of figure-ground influences were preserved. Effects of object knowledge and familiarity on figure-ground organization were also significantly diminished. Our results suggest that the mechanisms mediating contextual and familiarity influences on figure-ground organization are dissociable from those mediating local influences on figure-ground assignment. The disruption of contextual processing in intermediate visual areas may play a role in the substantial object recognition difficulties experienced by LG.

Counterbalancing for serial order carryover effects in experimental condition orders.

Reactions of neural, psychological, and social systems are rarely, if ever, independent of previous inputs and states. The potential for serial order carryover effects from one condition to the next in a sequence of experimental trials makes counterbalancing of condition order an essential part of experimental design. Here, a method is proposed for generating counterbalanced sequences for repeated-measures designs including those with multiple observations of each condition on one participant and self-adjacencies of conditions. Condition ordering is reframed as a graph theory problem. Experimental conditions are represented as vertices in a graph and directed edges between them represent temporal relationships between conditions. A counterbalanced trial order results from traversing an Euler circuit through such a graph in which each edge is traversed exactly once. This method can be generalized to counterbalance for higher order serial order carryover effects as well as to create intentional serial order biases. Modern graph theory provides tools for finding other types of paths through such graph representations, providing a tool for generating experimental condition sequences with useful properties.

Shifting attention in viewer- and object-based reference frames after unilateral brain injury.

The aims of the present study were to investigate the respective roles that object- and viewer-based reference frames play in reorienting visual attention, and to assess their influence after unilateral brain injury. To do so, we studied 16 right hemisphere injured (RHI) and 13 left hemisphere injured (LHI) patients. We used a cueing design that manipulates the location of cues and targets relative to a display comprised of two rectangles (i.e., objects). Unlike previous studies with patients, we presented all cues at midline rather than in the left or right visual fields. Thus, in the critical conditions in which targets were presented laterally, reorienting of attention was always from a midline cue. Performance was measured for lateralized target detection as a function of viewer-based (contra- and ipsilesional sides) and object-based (requiring reorienting within or between objects) reference frames. As expected, contralesional detection was slower than ipsilesional detection for the patients. More importantly, objects influenced target detection differently in the contralesional and ipsilesional fields. Contralesionally, reorienting to a target within the cued object took longer than reorienting to a target in the same location but in the uncued object. This finding is consistent with object-based neglect. Ipsilesionally, the means were in the opposite direction. Furthermore, no significant difference was found in object-based influences between the patient groups (RHI vs. LHI). These findings are discussed in the context of reference frames used in reorienting attention for target detection.

Cue competition affects temporal dynamics of edge-assignment in human visual cortex.

Edge-assignment determines the perception of relative depth across an edge and the shape of the closer side. Many cues determine edge-assignment, but relatively little is known about the neural mechanisms involved in combining these cues. Here, we manipulated extremal edge and attention cues to bias edge-assignment such that these two cues either cooperated or competed. To index their neural representations, we flickered figure and ground regions at different frequencies and measured the corresponding steady-state visual-evoked potentials (SSVEPs). Figural regions had stronger SSVEP responses than ground regions, independent of whether they were attended or unattended. In addition, competition and cooperation between the two edge-assignment cues significantly affected the temporal dynamics of edge-assignment processes. The figural SSVEP response peaked earlier when the cues causing it cooperated than when they competed, but sustained edge-assignment effects were equivalent for cooperating and competing cues, consistent with a winner-take-all outcome. These results provide physiological evidence that figure-ground organization involves competitive processes that can affect the latency of figural assignment.

Grouping puts figure-ground assignment in context by constraining propagation of edge assignment.

Figure-ground organization involves the assignment of edges to a figural shape on one or the other side of each dividing edge. Established visual cues for edge assignment primarily concern relatively local rather than contextual factors. In the present article, we show that an assignment for a locally unbiased edge can be affected by an assignment of a remote contextual edge that has its own locally biased assignment. We find that such propagation of edge assignment from the biased remote context occurs only when the biased and unbiased edges are grouped. This new principle, whereby grouping constrains the propagation of figural edge assignment, emerges from both subjective reports and an objective short-term edge-matching task. It generalizes from moving displays involving grouping by common fate and collinearity, to static displays with grouping by similarity of edge-contrast polarity, or apparent occlusion. Our results identify a new contextual influence on edge assignment. They also identify a new mechanistic relation between grouping and figure-ground processes, whereby grouping between remote elements can constrain the propagation of edge assignment between those elements. Supplemental materials for this article may be downloaded from http://app.psychonomic-journals.org/content/supplemental.

Edge-region grouping in figure-ground organization and depth perception.

Edge-region grouping (ERG) is proposed as a unifying and previously unrecognized class of relational information that influences figure-ground organization and perceived depth across an edge. ERG occurs when the edge between two regions is differentially grouped with one region based on classic principles of similarity grouping. The ERG hypothesis predicts that the grouped side will tend to be perceived as the closer, figural region. Six experiments are reported that test the predictions of the ERG hypothesis for 6 similarity-based factors: common fate, blur similarity, color similarity, orientation similarity, proximity, and flicker synchrony. All 6 factors produce the predicted effects, although to different degrees. In a 7th experiment, the strengths of these figural/depth effects were found to correlate highly with the strength of explicit grouping ratings of the same visual displays. The relations of ERG to prior results in the literature are discussed, and possible reasons for ERG-based figural/depth effects are considered. We argue that grouping processes mediate at least some of the effects we report here, although ecological explanations are also likely to be relevant in the majority of cases.

Visual hemispatial neglect, re-assessed.

Increased computer use in clinical settings offers an opportunity to develop new neuropsychological tests that exploit the control computers have over stimulus dimensions and timing. However, before adopting new tools, empirical validation is necessary. In the current study, our aims were twofold: to describe a computerized adaptive procedure with broad potential for neuropsychological investigations, and to demonstrate its implementation in testing for visual hemispatial neglect. Visual search results from adaptive psychophysical procedures are reported from 12 healthy individuals and 23 individuals with unilateral brain injury. Healthy individuals reveal spatially symmetric performance on adaptive search measures. In patients, psychophysical outcomes (as well as those from standard paper-and-pencil search tasks) reveal visual hemispatial neglect. Consistent with previous empirical studies of hemispatial neglect, lateralized impairments in adaptive conjunction search are greater than in adaptive feature search tasks. Furthermore, those with right hemisphere damage show greater lateralized deficits in conjunction search than do those with left hemisphere damage. We argue that adaptive tests, which automatically adjust to each individual's performance level, are efficient methods for both clinical evaluations and neuropsychological investigations and have the potential to detect subtle deficits even in chronic stages, when flagrant clinical signs have frequently resolved.

The occlusion illusion: partial modal completion or apparent distance?

In the occlusion illusion, the visible portion of a partly occluded object (eg a semicircle partly hidden behind a rectangle) appears to be significantly larger than a physically identical region that is fully visible. This illusion may occur either because the visual system 'fills in' a thin strip along the occluded border (the partial-modal-completion hypothesis) or because the partly occluded object is perceived as farther away (the apparent-distance hypothesis). We measured the magnitude of the occlusion illusion psychophysically in several experiments to investigate its causes. The results of experiments 1-3 are consistent with the general proposal that the magnitude of the illusion varies with the strength of the evidence for occlusion, supporting the inference that it is due to occlusion. Experiment 4 provides a critical test between apparent-distance and partial-modal-completion explanations by determining whether the increase in apparent size of the occluded region results from a change in its perceived shape (due to the modal extension of the occluded shape along the occluding edge, as predicted by the partial-modal-completion hypothesis) or from a change in its perceived overall size (as predicted by the apparent-distance hypothesis). The results more strongly support the partial-modal-completion hypothesis.

Crossing the midline: reducing attentional deficits via interhemispheric interactions.

Patients with unilateral neglect and extinction show a profound lack of awareness of stimuli presented contralateral to their lesion. However, many processes of perception are intact and contralesional stimuli seem to reach a high level of representation, perceptual and semantic. Some of these processes can work to decrease the magnitude of the attentional deficit. Here, we examine two of these intact processes, feature detection and perceptual grouping. First, we demonstrate that feature detection occurs in parallel in the contralesional visual fields of neglect and extinction patients. Second, we attempt to dissociate the influence of perceptual contours across the vertical meridian from the presence of an object or higher-level perceptual unit (or group) that may be created by these contours. We find that connections across the midline affect attentional deficits independently of the objects they may create. This suggests that several effects of grouping on neglect and extinction may be mediated by long-range cortical interactions that arise from connections across the vertical meridian.

When does grouping happen?

Recent research on perceptual grouping is described with particular emphasis on identifying the level(s) at which grouping factors operate. Contrary to the classical view of grouping as an early, two-dimensional, image-based process, recent experimental results show that it is strongly influenced by phenomena related to perceptual constancy, such as binocular depth perception, lightness constancy, amodal completion, and illusory contours. These findings imply that at least some grouping processes operate at the level of phenomenal perception rather than at the level of the retinal image. Preliminary evidence is reported showing that grouping can affect perceptual constancy, suggesting that grouping processes must also operate at an early, preconstancy level. If so, grouping may be a ubiquitous, ongoing aspect of visual organization that occurs for each level of representation rather than as a single stage that can be definitively localized relative to other perceptual processes.