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A popular model for sensory processing, known as predictive coding, proposes that incoming signals are iteratively compared with top-down predictions along a hierarchical processing scheme. At each step, error signals arising from differences between actual input and prediction are forwarded and recurrently minimized by updating internal models to finally be "explained away". However, the neuronal mechanisms underlying such computations and their limitations in processing speed are largely unknown. Further, it remains unclear at which step of cortical processing prediction errors are explained away, if at all. In the present study, human subjects briefly viewed the superposition of two orthogonally oriented gratings followed by abrupt removal of one orientation after either 33 or 200 milliseconds. Instead of strictly seeing the remaining orientation, observers report rarely but highly significantly an illusory percept of the arithmetic difference between previous and actual orientations. Previous findings in cats using the identical paradigm suggest that such difference signals are inherited from first steps of visual cortical processing. In light of early modeling accounts of predictive coding, in which visual neurons were interpreted as residual error detectors signaling the difference between actual input and its temporal prediction based on past input, our data may indicate continued access to residual errors. Such strategy permits time-critical perceptual decision making across a spectrum of competing internal signals up to the highest levels of processing. Thus, the occasional appearance of a prediction error-like illusory percept may uncover maintained flexibility at perceptual decision stages when subjects cope with highly dynamic and ambiguous visual stimuli.

During the observation of an ambiguous figure our perception alternates between mutually exclusive interpretations, although the stimulus itself remains unchanged. The rate of these endogenous reversals has been discussed as reflecting basic aspects of endogenous brain dynamics. Recent evidence indicates that extensive meditation practice evokes long-term functional and anatomic changes in the brain, also affecting the endogenous brain dynamics. As one of several consequences the rate of perceptual reversals during ambiguous figure perception decreases. In the present study we compared EEG-correlates of endogenous reversals of ambiguous figures between meditators and non-meditating controls in order to better understand timing and brain locations of this altered endogenous brain dynamics. A well-established EEG paradigm was used to measure the neural processes underlying endogenous perceptual reversals of ambiguous figures with high temporal precision. We compared reversal-related ERPs between experienced meditators and non-meditating controls. For both groups we found highly similar chains of reversal-related ERPs, starting early in visual areas, therewith replicating previous findings from the literature. Meditators, however, showed an additional frontal ERP signature already 160 ms after stimulus onset (Frontal Negativity). We interpret the additional, meditation-specific ERP results as evidence that extensive meditation practice provides control of frontal brain areas over early sensory processing steps. This may allow meditators to overcome phylogenetically evolved perceptual and attentional processing automatisms.

AIM: The present study utilizes perceptual hysteresis effects to compare the ambiguity of Mona Lisa's emotional face expression (high-level ambiguity) and of geometric cube stimuli (low-level ambiguity). METHODS: In two experiments we presented series of nine Mona Lisa variants and nine cube variants. Stimulus ambiguity was manipulated by changing Mona Lisa's mouth curvature (Exp. 1) and the cubes' back-layer luminance (Exp. 2). Each experiment consisted of three conditions, two with opposite stimulus presentation sequences with increasing and decreasing degrees of ambiguity, respectively, and a third condition with a random presentation sequence. Participants indicated happy or sad face percepts (Exp. 1) and alternative 3D cube percepts (Exp. 2) by key presses. We studied the influences of a priori perceptual biases (long-term memory) and presentation order (short-term memory) on perception. RESULTS: Perception followed sigmoidal functions of the stimulus ambiguity morphing parameters. The morphing parameter for the functions' inflection points depended strongly on stimulus presentation order with similar effect sizes but different signs for the two stimulus types (positive hysteresis / priming for the cubes; negative hysteresis / adaptation for Mona Lisa). In the random conditions, the inflection points were located in the middle between those from the two directional conditions for the Mona Lisa stimuli. For the cube stimuli, they were superimposed on one sigmoidal function for the ordered condition. DISCUSSION: The hysteresis effects reflect the influence of short-term memory during the perceptual disambiguation of ambiguous sensory information. The effects for the two stimulus types are of similar size, explaining up to 34% of the perceptual variance introduced by the paradigm. We explain the qualitative difference between positive and negative hysteresis with adaptation for Mona Lisa and with priming for the cubes. In addition, the hysteresis paradigm allows a quantitative determination of the impact of adaptation and priming during the resolution of perceptual ambiguities. The asymmetric shifts of inflection points in the case of the cube stimuli is likely due to an a priori perceptual bias, reflecting an influence of long-term memory. Whether corresponding influences also exist for the Mona Lisa variants is so far unclear.

Introduction: Consolidation is defined as the time necessary for memory stabilization after learning. In the present study we focused on effects of interference during the first 12 consolidation minutes after learning. Participants had to learn a set of German - Japanese word pairs in an initial learning task and a different set of German - Japanese word pairs in a subsequent interference task. The interference task started in different experimental conditions at different time points (0, 3, 6, and 9 min) after the learning task and was followed by subsequent cued recall tests. In a control experiment the interference periods were replaced by rest periods without any interference. Results: The interference task decreased memory performance by up to 20%, with negative effects at all interference time points and large variability between participants concerning both the time point and the size of maximal interference. Further, fast learners seem to be more affected by interference than slow learners. Discussion: Our results indicate that the first 12 min after learning are highly important for memory consolidation, without a general pattern concerning the precise time point of maximal interference across individuals. This finding raises doubts about the generalized learning recipes and calls for individuality of learning schedules.

The Necker-Zeno model of bistable perception provides a formal relation between the average duration of meta-stable percepts (dwell times T) of ambiguous figures and two other basic time scales (t(0), ΔT) underlying cognitive processing. The model predicts that dwell times T covary with t(0), ΔT or both. We tested this prediction by exploiting that observers, in particular experienced meditators, can volitionally control dwell times T. Meditators and non-meditators observed bistable Necker cubes either passively or tried to hold their current percept. The latencies of a centro-parietal event-related potential (CPP) were recorded as a physiological correlate of t(0). Dwell times T and the CPP latencies, correlated with t(0), differed between conditions and observer groups, while ΔT remained constant in the range predicted by the model. The covariation of CPP latencies and dwell times, as well as their quadratic functional dependence extends previous psychophysical confirmation of the Necker-Zeno model to psychophysiological measures.

The worldwide fascination of da Vinci's Mona Lisa has been dedicated to the emotional ambiguity of her face expression. In the present study we manipulated Mona Lisa's mouth curvature as one potential source of ambiguity and studied how a range of happier and sadder face variants influences perception. In two experimental conditions we presented different stimulus ranges with different step sizes between stimuli along the happy-sad axis of emotional face expressions. Stimuli were presented in random order and participants indicated the perceived emotional face expression (first task) and the confidence of their response (second task). The probability of responding 'happy' to the original Mona Lisa was close to 100%. Furthermore, in both conditions the perceived happiness of Mona Lisa variants described sigmoidal functions of the mouth curvature. Participants' confidence was weakest around the sigmoidal inflection points. Remarkably, the sigmoidal functions, as well as confidence values and reaction times, differed significantly between experimental conditions. Finally, participants responded generally faster to happy than to sad faces. Overall, the original Mona Lisa seems to be less ambiguous than expected. However, perception of and reaction to the emotional face content is relative and strongly depends on the used stimulus range.

BACKGROUND: During observation of the Necker cube perception becomes unstable and alternates repeatedly between a from-above-perspective ("fap") and a from-below-perspective ("fbp") interpretation. Both interpretations are physically equally plausible, however, observers usually show an a priori top-down bias in favor of the fap interpretation. Patients with Autism spectrum disorder are known to show an altered pattern of perception with a focus on sensory details. In the present study we tested whether this altered perceptual processing affects their reversal dynamics and reduces the perceptual bias during Necker cube observation. METHODS: 19 participants with Asperger syndrome and 16 healthy controls observed a Necker cube stimulus continuously for 5 minutes and indicated perceptual reversals by key press. We compared reversal rates (number of reversals per minute) and the distributions of dwell times for the two interpretations between observer groups. RESULTS: Asperger participants showed less perceptual reversal than controls. Six Asperger participants did not perceive any reversal at all, whereas all observers from the control group perceived at least five reversals within the five minutes observation time. Further, control participants showed the typical perceptual bias with significant longer median dwell times for the fap compared to the fbp interpretation. No such perceptual bias was found in the Asperger group. DISCUSSION: The perceptual system weights the incomplete and ambiguous sensory input with memorized concepts in order to construct stable and reliable percepts. In the case of the Necker cube stimulus, two perceptual interpretations are equally compatible with the sensory information and internal fluctuations may cause perceptual alternations between them-with a slightly larger probability value for the fap interpretation (perceptual bias). Smaller reversal rates in Asperger observers may result from the dominance of bottom-up sensory input over endogenous top-down factors. The latter may also explain the absence of a fap bias.

Environmental information available to our senses is incomplete and to varying degrees ambiguous. It has to be disambiguated in order to construct stable and reliable percepts. Ambiguous figures are artificial examples where perception is maximally unstable and alternates between possible interpretations. Tiny low-level changes can disambiguate an ambiguous figure and thus stabilize percepts. The present study compares ERPs evoked by ambiguous stimuli and disambiguated stimulus variants across three visual categories: geometry (Necker cube), motion (stroboscopic alternative motion stimulus, SAM) and semantics (Boring's old/young woman). We found that (a) disambiguated stimulus variants cause stable percepts and evoke two huge positive ERP excursions (Cohen's effect sizes 1-2), (b) the amplitudes of these ERP effects are inversely related to the degree of stimulus ambiguity, and (c) this pattern of results is consistent across all three tested visual categories. This generality across visual categories points to mechanisms at a very abstract (cognitive) level of processing. We discuss our results in the context of a high-level Bayesian inference unit that evaluates the reliability of perceptual processing results, given a priori incomplete, ambiguous sensory information. The ERP components may reflect the outcome of this reliability estimation.

Patients with schizophrenia have often been described as insensitive to nociceptive signals, but objective evidence is sparse. We address this question by combining subjective behavioral and objective neurochemical and neurophysiological measures. The present study involved 21 stabilized and mildly symptomatic patients with schizophrenia and 21 control subjects. We applied electrical stimulations below the pain threshold and assessed sensations of pain and unpleasantness with rating scales, and Somatosensory Evoked Potentials (SEPs/EEG). We also measured attention, two neurochemical stress indices (ACTH/cortisol), and subjective VEPs/EEG responses to visual emotional stimuli. Our results revealed that, subjectively, patients' evaluations do not differ from controls. However, the amplitude of EEG evoked potentials was greater in patients than controls as early as 50 ms after electrical stimulations and beyond one second after visual processing of emotional pictures. Such responses could not be linked to the stress induced by the stimulations, since stress hormone levels were stable. Nor was there a difference between patients and controls in respect of attention performance and tactile sensitivity. Taken together, all indices measured in patients in our study were either heightened or equivalent relative to healthy volunteers.

BACKGROUND: In von Schiller's Stroboscopic Alternative Motion (SAM) stimulus two visually presented diagonal dot pairs, located on the corners of an imaginary rectangle, alternate with each other and induce either horizontal, vertical or, rarely, rotational motion percepts. SAM motion perception can be described by a psychometric function of the dot aspect ratio ("AR", i.e. the relation between vertical and horizontal dot distances). Further, with equal horizontal and vertical dot distances (AR = 1) perception is biased towards vertical motion. In a series of five experiments, we presented tactile SAM versions and studied the role of AR and of different reference frames for the perception of tactile apparent motion. METHODS: We presented tactile SAM stimuli and varied the ARs, while participants reported the perceived motion directions. Pairs of vibration stimulators were attached to the participants' forearms and stimulator distances were varied within and between forearms. We compared straight and rotated forearm conditions with each other in order to disentangle the roles of exogenous and endogenous reference frames. RESULTS: Increasing the tactile SAM's AR biased perception towards vertical motion, but the effect was weak compared to the visual modality. We found no horizontal disambiguation, even for very small tactile ARs. A forearm rotation by 90° kept the vertical bias, even though it was now coupled with small ARs. A 45° rotation condition with crossed forearms, however, evoked a strong horizontal motion bias. DISCUSSION: Existing approaches to explain the visual SAM bias fail to explain the current tactile results. Particularly puzzling is the strong horizontal bias in the crossed-forearm conditions. In the case of tactile apparent motion, there seem to be no fixed priority rule for perceptual disambiguation. Rather the weighting of available evidence seems to depend on the degree of stimulus ambiguity, the current situation and on the perceptual strategy of the individual observer.

BACKGROUND: Asperger Autism is a lifelong psychiatric condition with highly circumscribed interests and routines, problems in social cognition, verbal and nonverbal communication, and also perceptual abnormalities with sensory hypersensitivity. To objectify both lower-level visual and cognitive alterations we looked for differences in visual event-related potentials (EEG) between Asperger observers and matched controls while they observed simple checkerboard stimuli. METHODS: In a balanced oddball paradigm checkerboards of two checksizes (0.6° and 1.2°) were presented with different frequencies. Participants counted the occurrence times of the rare fine or rare coarse checkerboards in different experimental conditions. We focused on early visual ERP differences as a function of checkerboard size and the classical P3b ERP component as an indicator of cognitive processing. RESULTS: We found an early (100-200 ms after stimulus onset) occipital ERP effect of checkerboard size (dominant spatial frequency). This effect was weaker in the Asperger than in the control observers. Further a typical parietal/central oddball-P3b occurred at 500 ms with the rare checkerboards. The P3b showed a right-hemispheric lateralization, which was more prominent in Asperger than in control observers. DISCUSSION: The difference in the early occipital ERP effect between the two groups may be a physiological marker of differences in the processing of small visual details in Asperger observers compared to normal controls. The stronger lateralization of the P3b in Asperger observers may indicate a stronger involvement of the right-hemispheric network of bottom-up attention. The lateralization of the P3b signal might be a compensatory consequence of the compromised early checksize effect. Higher-level analytical information processing units may need to compensate for difficulties in low-level signal analysis.

Ambiguous figures attract observers because perception alternates between different interpretations while the sensory information stays unchanged. Understanding the underlying processes is difficult because the precise time instant of this endogenous reversal event needs to be known but is difficult to measure. Presenting ambiguous figures discontinuously and using stimulus onset as estimation of the reversal event increased temporal resolution and provided a series of well-confirmed EEG signatures. In the current EEG study we used this 'onset paradigm' for the first time with Boring's old/young woman stimulus. We found an early occipital event-related potential (ERP) correlate of reversals between the perception of the old woman and the perception of the young woman that fits well with previous ERP findings. This component was not followed by the often-reported occipito-parietal Reversal Negativity at 260 ms, but instead by an occipito-temporal N170, that is typically reported with face stimuli. We interpret our results as follows: ambiguity conflicts take place during processing of stimulus elements in early visual areas roughly 130 ms after stimulus onset. The disambiguation of these elements and their assembly to object 'gestalts' result from an interplay between early visual and object-specific brain areas in a temporal window between 130 and 260 ms after stimulus onset. In the particular case of Boring's old/young woman the processes of element disambiguation and gestalt construction are already finished at 170 ms and, thus, 90 ms earlier than in the case of ambiguous geometric figures (eg Necker cube or Schroeder staircase) or of binocular rivalrous gratings.

Temporally distributed ("spaced") learning can be twice as efficient as massed learning. This "spacing effect" occurs with a broad spectrum of learning materials, with humans of different ages, with non-human vertebrates and also invertebrates. This indicates, that very basic learning mechanisms are at work ("generality"). Although most studies so far focused on very narrow spacing interval ranges, there is some evidence for a non-monotonic behavior of this "spacing effect" ("nonlinearity") with optimal spacing intervals at different time scales. In the current study we focused both the nonlinearity aspect by using a broad range of spacing intervals and the generality aspect by using very different learning/practice domains: Participants learned German-Japanese word pairs and performed visual acuity tests. For each of six groups we used a different spacing interval between learning/practice units from 7 min to 24 h in logarithmic steps. Memory retention was studied in three consecutive final tests, one, seven and 28 days after the final learning unit. For both the vocabulary learning and visual acuity performance we found a highly significant effect of the factor spacing interval on the final test performance. In the 12 h-spacing-group about 85% of the learned words stayed in memory and nearly all of the visual acuity gain was preserved. In the 24 h-spacing-group, in contrast, only about 33% of the learned words were retained and the visual acuity gain dropped to zero. The very similar patterns of results from the two very different learning/practice domains point to similar underlying mechanisms. Further, our results indicate spacing in the range of 12 hours as optimal. A second peak may be around a spacing interval of 20 min but here the data are less clear. We discuss relations between our results and basic learning at the neuronal level.

PURPOSE: We sought brain activity that predicts visual consciousness. METHODS: We used electroencephalography (EEG) to measure brain activity to a 1000-ms display of sine-wave gratings, oriented vertically in one eye and horizontally in the other. This display yields binocular rivalry: irregular alternations in visual consciousness between the images viewed by the eyes. We replaced both gratings with 200 ms of darkness, the gap, before showing a second display of the same rival gratings for another 1000 ms. We followed this by a 1000-ms mask then a 2000-ms inter-trial interval (ITI). Eleven participants pressed keys after the second display in numerous trials to say whether the orientation of the visible grating changed from before to after the gap or not. Each participant also responded to numerous non-rivalry trials in which the gratings had identical orientations for the two eyes and for which the orientation of both either changed physically after the gap or did not. RESULTS: We found that greater activity from lateral occipital-parietal-temporal areas about 180 ms after initial onset of rival stimuli predicted a change in visual consciousness more than 1000 ms later, on re-presentation of the rival stimuli. We also found that less activity from parietal, central, and frontal electrodes about 400 ms after initial onset of rival stimuli predicted a change in visual consciousness about 800 ms later, on re-presentation of the rival stimuli. There was no such predictive activity when the change in visual consciousness occurred because the stimuli changed physically. CONCLUSION: We found early EEG activity that predicted later visual consciousness. Predictive activity 180 ms after onset of the first display may reflect adaption of the neurons mediating visual consciousness in our displays. Predictive activity 400 ms after onset of the first display may reflect a less-reliable brain state mediating visual consciousness.

During observation of ambiguous figures our perception reverses spontaneously although the visual information stays unchanged. Research on this phenomenon so far suffered from the difficulty to determine the instant of the endogenous reversals with sufficient temporal precision. A novel experimental paradigm with discontinuous stimulus presentation improved on previous temporal estimates of the reversal event by a factor of three. It revealed that disambiguation of ambiguous visual information takes roughly 50 ms or two loops of recurrent neural activity. Further, the decision about the perceptual outcome has taken place at least 340 ms before the observer is able to indicate the consciously perceived reversal manually. We provide a short review about physiological studies on multistable perception with a focus on electrophysiological data. We further present a new perspective on multistable perception that can easily integrate previous apparently contradicting explanatory approaches. Finally we propose possible extensions toward other research fields where ambiguous figure perception may be useful as an investigative tool.

Repeated learning improves memory. Temporally distributed ("spaced") learning can be twice as efficient than massed learning. Importantly, learning success is a non-monotonic maximum function of the spacing interval between learning units. Further optimal spacing intervals seem to exist at different time scales from seconds to days. We briefly review the current state of knowledge about this "spacing effect" and then discuss very similar but so far little noticed spacing patterns during a form of synaptic plasticity at the cellular level, called long term potentiation (LTP). The optimization of learning is highly relevant for all of us. It may be realized easily with appropriate spacing. In our view, the generality of the spacing effect points to basic mechanisms worth for coordinated research on the different levels of complexity.

During observation of an ambiguous Necker cube, our percept changes spontaneously although the external stimulus does not. An EEG paradigm allowing time-resolved EEG measurement during endogenous perceptual reversals recently revealed a chain of ERP correlates beginning with an early occipital positivity at around 130 ms (Reversal Positivity, "RP"). In order to better understand the functional role of this RP, we investigated its relation to the P100, which is spatiotemporally close, typically occurring 100 ms after onset of a visual stimulus at occipital electrodes. We compared the relation of the ERP amplitudes to varying sizes of ambiguous Necker cubes. The main results are: (1) The P100 amplitude increases monotonically with stimulus size but is independent of the participants' percept. (2) The RP, in contrast, is percept-related and largely unaffected by stimulus size. (3) A similar pattern to RP was found for reaction times: They depend on the percept but not on stimulus size. We speculate that the P100 reflects processing of elementary visual features, while the RP is related to a processing conflict during 3D interpretation that precedes a reversal. The present results indicate that low-level visual processing (related to stimulus size) and (relative) high-level processing (related to perceptual reversal) occur in close spatial and temporal vicinity.

Ambiguous figures induce sudden transitions between rivaling percepts. We investigated electroencephalogram frequency modulations of accompanying change-related de- and rebinding processes. Presenting the stimuli discontinously, we synchronized perceptual reversals with stimulus onset, which served as a time reference for averaging. The resultant gain in temporal resolution revealed a sequence of time-frequency correlates of the reversal process. Most conspicuous was a transient right-hemispheric gamma modulation preceding endogenous reversals by at least 200 ms. No such modulation occurred with exogenously induced reversals of unambiguous stimulus variants. Post-onset components were delayed for ambiguous compared to unambiguous stimuli. The time course of oscillatory activity differed in several respects from predictions based on binding-related hypotheses. The gamma modulation preceding endogenous reversals may indicate an unstable brain state, ready to switch.