A phase-shifting anterior-posterior network organizes global phase relations.

Prior research has identified a variety of task-dependent networks that form through inter-regional phase-locking of oscillatory activity that are neural correlates of specific behaviors. Despite ample knowledge of task-specific functional networks, general rules governing global phase relations have not been investigated. To discover such general rules, we focused on phase modularity, measured as the degree to which global phase relations in EEG comprised distinct synchronized clusters interacting with one another at large phase lags. Synchronized clusters were detected with a standard community-detection algorithm, and the degree of phase modularity was quantified by the index q. Notably, we found that the mechanism controlling phase modularity is remarkably simple. A network comprising anterior-posterior long-distance connectivity coherently shifted phase relations from low-angles (|Δθ| < π/4) in low-modularity states (bottom 5% in q) to high-angles (|Δθ| > 3π/4) in high-modularity states (top 5% in q), accounting for fluctuations in phase modularity. This anterior-posterior network may play a fundamental functional role as (1) it controls phase modularity across a broad range of frequencies (3-50 Hz examined) in different behavioral conditions (resting with the eyes closed or watching a silent nature video) and (2) neural interactions (measured as power correlations) in beta-to-gamma bands were consistently elevated in high-modularity states. These results may motivate future investigations into the functional roles of phase modularity as well as the anterior-posterior network that controls it.

Radial bias alters high-level motion perception.

The visual system involves various orientation and visual field anisotropies, one of which is a preference for radial orientations and motion directions. By radial, we mean those directions coursing symmetrically outward from the fovea into the periphery. This bias stems from anatomical and physiological substrates in the early visual system. We recently reported that this low-level visual anisotropy can alter perceived object orientation. Here, we report that radial bias can also alter another higher-level system, the perceived direction of apparent motion. We presented a bistable apparent motion quartet in the center of the screen while participants fixated on various locations around the quartet. Participants (N = 22) were strongly biased to see the motion direction that was radial with respect to their fixation, controlling for any biases with center fixation. This was observed using a vertical-horizontal quartet as well as an oblique quartet (45° rotated quartet). The latter allowed us to rule out the contribution of the hemisphere effect where motion across the midline is perceived less often. These results extend our earlier findings on perceived object orientation, showing that low-level structural aspects of the visual system alter yet another higher-level visual process, that of apparent motion perception.

Motion duration is overestimated behind an occluder in action and perception tasks.

Motion estimation behind an occluder is a common task in situations like crossing the street or passing another car. People tend to overestimate the duration of an object's motion when it gets occluded for subsecond motion durations. Here, we explored (a) whether this bias depended on the type of interceptive action: discrete keypress versus continuous reach and (b) whether it was present in a perception task without an interceptive action. We used a prediction-motion task and presented a bar moving across the screen with a constant velocity that later became occluded. In the action task, participants stopped the occluded bar when they thought the bar reached the goal position via keypress or reach. They were more likely to stop the bar after it passed the goal position regardless of the action type, suggesting that the duration of occluded motion was overestimated (or its speed was underestimated). In the perception task, where participants judged whether a tone was presented before or after the bar reached the goal position, a similar bias was observed. In both tasks, the bias was near constant across motion durations and directions and grew over trials. We speculate that this robust bias may be due to a temporal illusion, Bayesian slow-motion prior, or the processing of the visible-occluded boundary crossing. Understanding its exact mechanism, the conditions on which it depends, and the relative roles of speed and time perception requires further research.

Radial Bias Alters Perceived Object Orientation.

Orientation sensitivity is a fundamental property of the visual system, but not all orientations are created equal. For instance, radially oriented stimuli, aligned with a line intersecting the center of gaze, produce greater activity throughout the visual cortex and are associated with greater perceptual sensitivity compared with other orientations. Here, we discuss a robust visual illusion that is likely related to this preference. Using a continuous response measure, participants (N = 36 adults) indicated the gap position in a peripheral Landolt C placed in one of eight orientations and eight locations along four meridians (vertical, horizontal, 45°, 135°). The error distributions revealed that the perceived gap was attracted toward the radial axis. For instance, the gap in a regular C would often be wrongly perceived as tilted 45° corresponding to the oblique meridian where it was placed. These findings demonstrate an unsuspected early-vision influence on the perceived orientation of an object.

Spatiotemporal dynamics of maximal and minimal EEG spectral power.

Oscillatory neural activities are prevalent in the brain with their phase realignment contributing to the coordination of neural communication. Phase realignments may have especially strong (or weak) impact when neural activities are strongly synchronized (or desynchronized) within the interacting populations. We report that the spatiotemporal dynamics of strong regional synchronization measured as maximal EEG spectral power-referred to as activation-and strong regional desynchronization measured as minimal EEG spectral power-referred to as suppression-are characterized by the spatial segregation of small-scale and large-scale networks. Specifically, small-scale spectral-power activations and suppressions involving only 2-7% (1-4 of 60) of EEG scalp sites were prolonged (relative to stochastic dynamics) and consistently co-localized in a frequency specific manner. For example, the small-scale networks for θ, α, ß1, and ß2 bands (4-30 Hz) consistently included frontal sites when the eyes were closed, whereas the small-scale network for γ band (31-55 Hz) consistently clustered in medial-central-posterior sites whether the eyes were open or closed. Large-scale activations and suppressions involving over 17-30% (10-18 of 60) of EEG sites were also prolonged and generally clustered in regions complementary to where small-scale activations and suppressions clustered. In contrast, intermediate-scale activations and suppressions (involving 7-17% of EEG sites) tended to follow stochastic dynamics and were less consistently localized. These results suggest that strong synchronizations and desynchronizations tend to occur in small-scale and large-scale networks that are spatially segregated and frequency specific. These synchronization networks may broadly segregate the relatively independent and highly cooperative oscillatory processes while phase realignments fine-tune the network configurations based on behavioral demands.

Probabilistic, entropy-maximizing control of large-scale neural synchronization.

Oscillatory neural activity is dynamically controlled to coordinate perceptual, attentional and cognitive processes. On the macroscopic scale, this control is reflected in the U-shaped deviations of EEG spectral-power dynamics from stochastic dynamics, characterized by disproportionately elevated occurrences of the lowest and highest ranges of power. To understand the mechanisms that generate these low- and high-power states, we fit a simple mathematical model of synchronization of oscillatory activity to human EEG data. The results consistently indicated that the majority (~95%) of synchronization dynamics is controlled by slowly adjusting the probability of synchronization while maintaining maximum entropy within the timescale of a few seconds. This strategy appears to be universal as the results generalized across oscillation frequencies, EEG current sources, and participants (N = 52) whether they rested with their eyes closed, rested with their eyes open in a darkened room, or viewed a silent nature video. Given that precisely coordinated behavior requires tightly controlled oscillatory dynamics, the current results suggest that the large-scale spatial synchronization of oscillatory activity is controlled by the relatively slow, entropy-maximizing adjustments of synchronization probability (demonstrated here) in combination with temporally precise phase adjustments (e.g., phase resetting generated by sensorimotor interactions). Interestingly, we observed a modest but consistent spatial pattern of deviations from the maximum-entropy rule, potentially suggesting that the mid-central-posterior region serves as an "entropy dump" to facilitate the temporally precise control of spectral-power dynamics in the surrounding regions.

Revealing the effects of temporal orienting of attention on response conflict using continuous movements.

Orienting attention in time enables us to prepare for forthcoming perception and action (e.g., estimating the duration of a yellow traffic light when driving). While temporal orienting can facilitate performance on simple tasks, its influence on complex tasks involving response conflict is unclear. Here, we adapted the flanker paradigm to a choice-reaching task where participants used a computer mouse to reach to the left or right side of the screen, as indicated by the central arrow presented with either the congruent or incongruent flankers. We assessed the effects of temporal orienting by manipulating goal-driven temporal expectation (using probabilistic variations in target timing) and stimulus-driven temporal priming (using sequential repetitions versus switches in target timing). We tested how temporal orienting influenced the dynamics of response conflict resolution. Recent choice-reaching studies have indicated that under response conflict, delayed movement initiation captures the response threshold adjustment process, whereas increased curvature toward the incorrect response captures the degree of coactivation of the response alternatives during the controlled response selection process. Both temporal expectation and priming reduced the initiation latency regardless of response conflict, suggesting that both lowered response thresholds independently of response conflict. Notably, temporal expectation, but not temporal priming, increased the curvature toward the incorrect response on incongruent trials. These results suggest that temporal orienting generally increases motor preparedness, but goal-driven temporal orienting particularly interferes with response conflict resolution, likely through its influence on response thresholds. Overall, our study highlights the interplay between temporal orienting and cognitive control in goal-directed action.

EEG state-trajectory instability and speed reveal global rules of intrinsic spatiotemporal neural dynamics.

Spatiotemporal dynamics of EEG/MEG (electro-/magneto-encephalogram) have typically been investigated by applying time-frequency decomposition and examining amplitude-amplitude, phase-phase, or phase-amplitude associations between combinations of frequency bands and scalp sites, primarily to identify neural correlates of behaviors and traits. Instead, we directly extracted global EEG spatiotemporal dynamics as trajectories of k-dimensional state vectors (k = the number of estimated current sources) to investigate potential global rules governing neural dynamics. We chose timescale-dependent measures of trajectory instability (approximately the 2nd temporal derivative) and speed (approximately the 1st temporal derivative) as state variables, that succinctly characterized trajectory forms. We compared trajectories across posterior, central, anterior, and lateral scalp regions as the current sources under those regions may serve distinct functions. We recorded EEG while participants rested with their eyes closed (likely engaged in spontaneous thoughts) to investigate intrinsic neural dynamics. Some potential global rules emerged. Time-averaged trajectory instability from all five regions tightly converged (with their variability minimized) at the level of generating nearly unconstrained but slightly conservative turns (~100° on average) on the timescale of ~25 ms, suggesting that spectral-amplitude profiles are globally adjusted to maintain this convergence. Further, within-frequency and cross-frequency phase relations appear to be independently coordinated to reduce average trajectory speed and increase the variability in trajectory speed and instability in a relatively timescale-invariant manner, and to make trajectories less oscillatory. Future research may investigate the functional relevance of these intrinsic global-dynamics rules by examining how they adjust to various sensory environments and task demands or remain invariant. The current results also provide macroscopic constraints for quantitative modeling of neural dynamics as the timescale dependencies of trajectory instability and speed are relatable to oscillatory dynamics.

Rhythm Violation Enhances Auditory-Evoked Responses to the Extent of Overriding Sensory Adaptation in Passive Listening.

Sensory systems utilize temporal structure in the environment to build expectations about the timing of forthcoming events. We investigated the effects of rhythm-based temporal expectation on auditory responses measured with EEG recorded from the frontocentral sites implicated in auditory processing. By manipulating temporal expectation and the interonset interval (IOI) of tones, we examined how neural responses adapted to auditory rhythm and reacted to stimuli that violated the rhythm. Participants passively listened to the tones while watching a silent nature video. In Experiment 1 (n = 22), in the long-IOI block, tones were frequently presented (80%) with 1.7-sec IOI and infrequently presented (20%) with 1.2-sec IOI, generating unexpectedly early tones that violated temporal expectation. Conversely, in the short-IOI block, tones were frequently presented with 1.2-sec IOI and infrequently presented with 1.7-sec IOI, generating late tones. We analyzed the tone-evoked N1-P2 amplitude of ERPs and intertrial phase clustering in the theta-alpha band. The results provided evidence of strong delay-dependent adaptation effects (short-term, sensitive to IOI), weak cumulative adaptation effects (long-term, driven by tone repetition over time), and robust temporal-expectation violation effects over and above the adaptation effects. Experiment 2 (n = 22) repeated Experiment 1 with shorter IOIs of 1.2 and 0.7 sec. Overall, we found evidence of strong delay-dependent adaptation effects, weak cumulative adaptation effects (which may most efficiently accumulate at the tone presentation rate of ∼1 Hz), and robust temporal-expectation violation effects that substantially boost auditory responses to the extent of overriding the delay-dependent adaptation effects likely through mechanisms involved in exogenous attention.

Spectral-power associations reflect amplitude modulation and within-frequency interactions on the sub-second timescale and cross-frequency interactions on the seconds timescale.

We investigated the global structure of intrinsic cross-frequency dynamics by systematically examining power-based temporal associations among a broad range of oscillation frequencies both within and across EEG-based current sources (sites). We focused on power-based associations that could reveal unique timescale dependence independently of interacting frequencies. Large spectral-power fluctuations across all sites occurred at two characteristic timescales, sub-second and seconds, yielding distinct patterns of cross-frequency associations. On the fast sub-second timescale, within-site (local) associations were consistently between pairs of ß-γ frequencies differing by a constant Δf (particularly Δf ~ 10 Hz at posterior sites and Δf ~ 16 Hz at lateral sites) suggesting that higher-frequency oscillations are organized into Δf amplitude-modulated packets, whereas cross-site (long-distance) associations were all within-frequency (particularly in the >30 Hz and 6-12 Hz ranges, suggestive of feedforward and feedback interactions). On the slower seconds timescale, within-site (local) associations were characterized by a broad range of frequencies selectively associated with ~10 Hz at posterior sites and associations among higher (>20 Hz) frequencies at lateral sites, whereas cross-site (long-distance) associations were characterized by a broad range of frequencies at posterior sites selectively associated with ~10 Hz at other sites, associations among higher (>20 Hz) frequencies among lateral and anterior sites, and prevalent associations at ~10 Hz. Regardless of timescale, within-site (local) cross-frequency associations were weak at anterior sites indicative of frequency-specific operations. Overall, these results suggest that the fast sub-second-timescale coordination of spectral power is limited to local amplitude modulation and insulated within-frequency long-distance interactions (likely feedforward and feedback interactions), while characteristic patterns of cross-frequency interactions emerge on the slower seconds timescale. The results also suggest that the occipital α oscillations play a role in organizing higher-frequency oscillations into ~10 Hz amplitude-modulated packets to communicate with other regions. Functional implications of these timescale-dependent cross-frequency associations await future investigations.

Correction: Neural, functional, and aesthetic impacts of spatially heterogeneous flicker: A potential role of natural flicker.

[This corrects the article DOI: 10.1371/journal.pone.0219107.].

Neural, functional, and aesthetic impacts of spatially heterogeneous flicker: A potential role of natural flicker.

Spatially heterogeneous flicker, characterized by probabilistic and locally independent luminance modulations, abounds in nature. It is generated by flames, water surfaces, rustling leaves, and so on, and it is pleasant to the senses. It affords spatiotemporal multistability that allows sensory activation conforming to the biases of the visual system, thereby generating the perception of spontaneous motion and likely facilitating the calibration of motion detectors. One may thus hypothesize that spatially heterogeneous flicker might potentially provide restoring stimuli to the visual system that engage fluent (requiring minimal top-down control) and self-calibrating processes. Here, we present some converging behavioral and electrophysiological evidence consistent with this idea. Spatially heterogeneous (multistable) flicker (relative to controls matched in temporal statistics) reduced posterior EEG (electroencephalography) beta power implicated in long-range neural interactions that impose top-down influences on sensory processing. Further, the degree of spatiotemporal multistability, the amount of posterior beta-power reduction, and the aesthetic responses to flicker were closely associated. These results are consistent with the idea that the pleasantness of natural flicker may derive from its spatiotemporal multistability that affords fluent and self-calibrating visual processing.

Probability-driven and stimulus-driven orienting of attention to time and sensory modality.

The timing and the sensory modality of behaviorally relevant events often vary predictably, so that it is beneficial to adapt the sensory system to their statistical regularities. Indeed, statistical information about target timing and/or sensory modality modulates behavioral responses-called expectation effects. Responses are also facilitated by short-term repetitions of target timing and/or sensory modality-called priming effects. We examined how the expectation and priming effects on target timing (short vs. long cue-to-target interval) and target modality (auditory vs. visual) interacted. Temporal expectation was manipulated across blocks, while modality expectation was manipulated across participants. Responses were faster when targets were presented at the expected timing and/or in the expected modality in an additive manner, suggesting that temporal and modality expectation operate relatively independently. Similarly, responses were faster when the timing and/or modality of targets was repeated across trials in an additive manner, suggesting that temporal and modality priming operate relatively independently. Importantly, the interactions between expectation and priming were domain specific. In the temporal domain, temporal-expectation effects were observed only when temporal-priming effects were absent. In the modality domain, modality-priming effects predominated for auditory targets whereas modality-expectation effects predominated for visual targets. Thus, the interactions between probability-driven expectation and stimulus-driven priming processes appear to be controlled separately for the mechanisms that direct attention to specific temporal intervals and for the mechanisms that direct attention to specific sensory modalities. These results may suggest that the sensory system concurrently optimizes attentional priorities within temporal and sensory-modality domains.

Near complete interocular transfer of the attentional repulsion effect.

During a brief period following attention capture by an abrupt-onset cue, a briefly presented item in the vicinity appears to be displaced away from the focus of attention. This effect, termed the attentional repulsion effect (ARE), can be induced with various ways of focusing attention (e.g., color pop-out, an auditory cue, voluntary focusing), and can be measured in various ways (e.g., as a vernier offset, shape deformation, action error). While most prior results on ARE have confirmed its close relationship with attention mechanisms, DiGiacomo and Pratt Vision Research 64 (2012) 35-41 reported no interocular transfer of ARE, placing ARE's operational locus at the level of monocular processing in V1 and/or LGN. DiGiacomo's and Pratt's result is surprising because even local pattern adaptation effects thought to be mediated by V1 show 50%-80% of interocular transfer. How could it be that a strongly attention-dependent effect is exclusively mediated by monocular processes? It was thus important to replicate DiGiacomo's and Pratt's surprising results using a transient-free mirror-based stereoscope and a broader method where ARE was measured with both vertical and horizontal vernier offsets. Our results demonstrate a nearly complete interocular transfer of ARE, with stronger ARE obtained with horizontal than with vertical verniers, implying that ARE may be hemifield dependent. We speculate that the null ARE result reported by DiGiacomo and Pratt in their dichoptic condition may be due to a statistical anomaly or to a potential visual artifact generated by the eye shutters that were used to present dichoptic stimuli.

Temporal expectation weights visual signals over auditory signals.

Temporal expectation is a process by which people use temporally structured sensory information to explicitly or implicitly predict the onset and/or the duration of future events. Because timing plays a critical role in crossmodal interactions, we investigated how temporal expectation influenced auditory-visual interaction, using an auditory-visual crossmodal congruity effect as a measure of crossmodal interaction. For auditory identification, an incongruent visual stimulus produced stronger interference when the crossmodal stimulus was presented with an expected rather than an unexpected timing. In contrast, for visual identification, an incongruent auditory stimulus produced weaker interference when the crossmodal stimulus was presented with an expected rather than an unexpected timing. The fact that temporal expectation made visual distractors more potent and visual targets less susceptible to auditory interference suggests that temporal expectation increases the perceptual weight of visual signals.

Comparing the effects of implicit and explicit temporal expectation on choice response time and response conflict.

People can use temporally structured sensory information to anticipate future events. Temporal information can be presented implicitly through probability manipulation without participants' awareness of the manipulation, or explicitly conveyed through instructions. We examined how implicit and explicit temporal information established temporal expectations that influenced choice response times and response conflict (measured as flanker effects). We implicitly manipulated temporal structure by block-wise varying the likely timing of a target. In the short-interval block, a target was presented frequently (80 % of trials) after a short (400 ms) cue-to-target interval and infrequently (20 % of trials) after a long (1200 ms) interval; the probability assignment was reversed in the long-interval block. Building on this baseline condition (Experiment 1), we augmented the temporal information by filling the cue-to-target intervals with tones (Experiment 2), explicitly informed participants of the prevalent time interval (Experiment 3) and provided trial-by-trial reminders of the prevalent time interval (Experiment 4). The temporal probability manipulation alone (of which participants were unaware) influenced choice response times but only when the temporal information was augmented with tones, whereas providing the explicit knowledge of the temporal manipulation, with or without trial-by-trial reminders, robustly influenced choice response times. Response conflict was unaffected by these conditions. These results suggest that temporal expectation can be established by the implicit learning of a temporal structure given that sufficiently strong temporal information is presented as well as by the explicit knowledge of the temporal structure. This established temporal expectation influences choice response times without necessarily affecting the strength of response conflict.