Perceptual phenomena in destructured sensory fields: Probing the brain's intrinsic functional architectures.

Destructured sensory fields, involving homogenous stimulation with little or no time-varying structure, provide a fertile ground for testing hypotheses about predictive coding in the human brain. Extended exposure to sensory patterns that deviate substantially from the statistics of natural environments can elicit a bewildering range of perceptual phenomena, up to and including vivid oneiric imagery. We illustrate how this large variety of perceptual effects can be understood as the experiential counterpart of auto-generated neuronal dynamics, unconstrained by parameters that tune the waking sensorium. We synthesize the literature on autonomous neuronal activity across multiple spatiotemporal scales with generative models of brain function and evidence from artificial neural architectures. Perception, we argue, emerges from a process of non-random sampling from an intrinsic distribution of hypotheses rather than a direct transfer of information from the world. The imagery that occurs in altered sensory environments is explained as the outcome of an iterative search through internal world models in which the structural typology of percepts reflects the brain's intrinsic functional architectures.

Effects of emotional valence and arousal on acoustic duration reproduction assessed via the "dual klepsydra model".

We report results of an acoustic duration reproduction task with stimulus duration of 2, 4, and 6 s, using 45 emotionally negative, positive, and neutral sounds from the International Affective Digitized Sounds System, in a sample of 31 young healthy participants. To investigate the influence of induced emotions on perceived duration, the effects of emotional modulation were quantified in two ways: (1) via model-free indices (aggregated ratios of reproduced times), and (2) via dual klepsydra model (dkm)-based estimates of parameters of internal time representation. Both data-analytic approaches reveal an effect of emotional valence/arousal, namely, a significantly longer reproduction response for emotional stimuli than for the neutral stimuli. The advantage of the dkm-based approach is its ability to disentangle stimulus-related effects, which are represented by "flow intensities," from general effects which are due to the lossy character of temporal integration. We explain the rationale of the dkm-based strategy and interpret the observed effect within the dkm-framework as transient increase of internal "flows." This interpretation is in line with recent conceptualizations of an "embodiment" of time where the model-posited flows correspond to the ongoing stream of interoceptive (bodily) neural signals. Neurophysiological findings on correlations between the processing of body signals and the perception of time provide cumulative evidence for this working hypothesis.

Perception of acoustically presented time series with varied intervals.

Data from three experiments on serial perception of temporal intervals in the supra-second domain are reported. Sequences of short acoustic signals ("pips") separated by periods of silence were presented to the observers. Two types of time series, geometric or alternating, were used, where the modulus 1+δ of the inter-pip series and the base duration Tb (range from 1.1 to 6s) were varied as independent parameters. The observers had to judge whether the series were accelerating, decelerating, or uniform (3 paradigm), or to distinguish regular from irregular sequences (2 paradigm). "Intervals of subjective uniformity" (isus) were obtained by fitting Gaussian psychometric functions to individual subjects' responses. Progression towards longer base durations (Tb=4.4 or 6s) shifts the isus towards negative δs, i.e., accelerating series. This finding is compatible with the phenomenon of "subjective shortening" of past temporal intervals, which is naturally accounted for by the lossy integration model of internal time representation. The opposite effect observed for short durations (Tb=1.1 or 1.5s) remains unexplained by the lossy integration model, and presents a challenge for further research.

Duration reproduction: lossy integration and effects of sensory modalities, cognitive functioning, age, and sex.

The "dual klepsydra model" (DKM) of internal time representation successfully models duration reproduction data, but relations between the DKM-based parameter kappa ("loss rate") and procedural variables (presentation modality) or individual characteristics (cognitive indices, age, sex) remained as yet unexplored. For that purpose, were-analyzed data from an earlier time reproduction study (N = 100), using visually or acoustically presented intervals of 1-5 sec. duration. Typical values of parameter kappa were approximately 0.03-0.04 sec.(-1), corresponding to relaxation times of internal "lossy integrators" of approximately 30 sec. Significant effects of presentation modality (smaller kappa values for the visual reproduction task) and of age (greater kappa in acoustic reproduction with increasing age) were observed. Cognitive variables (working memory, general fluid reasoning, attention) and sex of participants were not associated with kappa. Cognitive functions seem to play only a minor, if any, role at the level of time representation addressed by the DKM.

Determinants of filled/empty optical illusion: differential effects of patterning.

A subdivided path in the visual field appears longer than an empty path of the same length. This effect may be attributed to the division of the path into multiple segments, or to an influence of the visual elements used to mark the subdivision, and thus filling-up the estimated space. To address this question, we used two series of stimuli, in which the spatial distribution of the filling optical mater, or the form of the dividers, was varied while the relative coverage of the filled space was kept constant. We found significant dependence of the effect magnitude on a number of filling elements as well as on their form. These results indicate that the illusory space expansion is not merely an effect of 'filling-up' the space, but it also depends on the filling pattern. Consequences of these findings for the theory of the Oppel-Kundt phenomenon are briefly discussed.

Abnormal activity in the precuneus during time perception in Parkinson's disease: an fMRI study.

BACKGROUND: Parkinson's disease (PD) patients are deficient in time estimation. This deficit improves after dopamine (DA) treatment and it has been associated with decreased internal timekeeper speed, disruption of executive function and memory retrieval dysfunction. METHODOLOGY/FINDINGS: The aim of the present study was to explore the neurophysiologic correlates of this deficit. We performed functional magnetic resonance imaging on twelve PD patients while they were performing a time reproduction task (TRT). The TRT consisted of an encoding phase (during which visual stimuli of durations from 5 s to 16.6 s, varied at 8 levels were presented) and a reproduction phase (during which interval durations were reproduced by a button pressing). Patients were scanned twice, once while on their DA medication (ON condition) and once after medication withdrawal (OFF condition). Differences in Blood-Oxygenation-Level-Dependent (BOLD) signal in ON and OFF conditions were evaluated. The time course of activation in the brain areas with different BOLD signal was plotted. There were no significant differences in the behavioral results, but a trend toward overestimation of intervals ≤11.9 s and underestimation of intervals ≥14.1 s in the OFF condition (p<0.088). During the reproduction phase, higher activation in the precuneus was found in the ON condition (p<0.05 corrected). Time course was plotted separately for long (≥14.1 s) and short (≤11.9 s) intervals. Results showed that there was a significant difference only in long intervals, when activity gradually decreased in the OFF, but remained stable in the ON condition. This difference in precuneus activation was not found during random button presses in a control task. CONCLUSIONS/SIGNIFICANCE: Our results show that differences in precuneus activation during retrieval of a remembered duration may underlie some aspects of time perception deficit in PD patients. We suggest that DA medication may allow compensatory activation in the precuneus, which results in a more accurate retrieval of remembered interval duration.

Determinants of filled/empty optical illusion: Influence of luminance contrast and polarity.

Subjective estimates of lengths or areas in the visual field depend on the visual contents of the estimated space (filled/empty or Oppel-Kundt illusion). We studied the dependence of this phenomenon on the presentation mode (white on black vs. black on white background), and on the figure/ground contrast. We found, as expected, overestimation of the filled part of the figure for both contrast polarities. The expansion effect was found to be an increasing function of the absolute luminance contrast, and was consistently higher for the negative (luminant figures on a dark background) than for the positive polarity. The contrast factor contributes from one-fifth to one-third of the total effect. Possible interpretations in terms of known sensory phenomena (irradiation, lateral interactions) or higher, integrative functions are discussed.

Theta-burst transcranial magnetic stimulation over the supplementary motor area decreases variability of temporal estimates.

OBJECTIVES: Supplementary motor area (SMA) was suggested to have a dominant role in the temporal control of behavior by many neuroimaging studies. The aim of this study was to support this hypothesis by influencing time estimates with theta burst transcranial magnetic stimulation (TMS) over the SMA. METHODS: Nineteen healthy volunteers with a mean age 25.9±3 (SD) years performed the time reproduction task (TRT) before and after 190 seconds of intermittent theta-burst TMS over SMA and the precuneus (total 600 pulses). The TRT consisted of an encoding phase (during which visual stimuli with durations of 5, 10 and 16.82 seconds were presented pseudorandomly) and a reproduction phase (during which interval durations were reproduced by pressing a button). Mean subjects' interval estimates as a measure of accuracy and standard deviation as a measure of variability pre-TMS and post-TMS were compared. RESULTS: Theta-burst TMS over both areas had no effect on the accuracy of duration estimates. An increased variability of interval reproduction was present after stimulation of the precuneus (p<0.01) with the biggest effect on the five second interval. Stimulation of SMA caused a decrease of variability in the ten second interval only (p<0.05). CONCLUSION: It is likely that increased variability of time estimates is a non-specific result of impaired attention and working memory after theta-burst TMS. Decreased variability after stimulation over the SMA could be explained in terms of enhanced activity of the physiological oscillator with a frequency close to 0.1 Hz.

Neural substrates of time perception and impulsivity.

Several studies provide empirical evidence for the association between impulsivity and time perception. However, little is known about the neural substrates underlying this function. This investigation examined the influence of impulsivity on neural activation patterns during the encoding and reproduction of intervals with durations of 3, 9 and 18s using event-related functional magnetic resonance imaging (fMRI). Twenty-seven subjects participated in this study, including 15 high impulsive subjects that were classified based on their self-rating. FMRI activation during the duration reproduction task was correlated with measures of two self-report questionnaires related to the concept of impulsivity (Barratt Impulsiveness Scale, BIS; Zimbardo Time Perspective Inventory, ZTPI). Behaviorally, those individuals who under-reproduced temporal intervals also showed lower scores on the ZTPI future perspective subscale and higher scores on the BIS. FMRI activation revealed an accumulating pattern of neural activity peaking at the end of the 9- and 18-s intervals within right posterior insula. Activations of brain regions during the reproduction phase of the timing task, such as those related to motor execution as well as to the 'core control network' - encompassing the inferior frontal and medial frontal cortices, the anterior insula as well as the inferior parietal cortex - were significantly correlated with reproduced duration, as well as with BIS and ZTPI subscales. In particular, the greater activation in these regions the shorter were the reproduced intervals, the more impulsive was an individual and the less pronounced the future perspective. Activation in the core control network, thus, may form a biological marker for cognitive time management and for impulsiveness.

Flicker-light induced visual phenomena: frequency dependence and specificity of whole percepts and percept features.

Flickering light induces visual hallucinations in human observers. Despite a long history of the phenomenon, little is known about the dependence of flicker-induced subjective impressions on the flicker frequency. We investigate this question using Ganzfeld stimulation and an experimental paradigm combining a continuous frequency scan (1-50 Hz) with a focus on re-occurring, whole percepts. On the single-subject level, we find a high degree of frequency stability of percepts. To generalize across subjects, we apply two rating systems, (1) a set of complex percept classes derived from subjects' reports and (2) an enumeration of elementary percept features, and determine distributions of occurrences over flicker frequency. We observe a stronger frequency specificity for complex percept classes than elementary percept features. Comparing the similarity relations among percept categories to those among frequency profiles, we observe that though percepts are preferentially induced by particular frequencies, the frequency does not unambiguously determine the experienced percept.

Genetic determinants of time perception mediated by the serotonergic system.

BACKGROUND: The present study investigates neurobiological underpinnings of individual differences in time perception. METHODOLOGY: Forty-four right-handed Russian Caucasian males (18-35 years old) participated in the experiment. The polymorphism of the genes related to the activity of serotonin (5-HT) and dopamine (DA)-systems (such as 5-HTT, 5HT2a, MAOA, DAT, DRD2, COMT) was determined upon the basis of DNA analysis according to a standard procedure. Time perception in the supra-second range (mean duration 4.8 s) was studied, using the duration discrimination task and parametric fitting of psychometric functions, resulting in individual determination of the point of subjective equality (PSE). Assuming the 'dual klepsydra model' of internal duration representation, the PSE values were transformed into equivalent values of the parameter κ (kappa), which is a measure of the 'loss rate' of the duration representation. An association between time representation parameters (PSE and κ, respectively) and 5-HT-related genes was found, but not with DA-related genes. Higher 'loss rate' (κ) of the cumulative duration representation were found for the carriers of genotypes characterized by higher 5-HT transmission, i.e., 1) lower 5-HT reuptake, known for the 5-HTTLPR SS polymorphism compared with LL, 2) lower 5-HT degradation, described for the 'low expression' variant of MAOA VNTR gene compared with 'high expression' variant, and 3) higher 5-HT2a receptor density, proposed for the TT polymorphism of 5-HT2a T102C gene compared with CC. CONCLUSION: Convergent findings of the present study and previous psychopharmacological studies suggest an action path from 5-HT-activity-related genes, via activity of 5-HT in the brain, to time perception. An involvement of the DA-system in the encoding of durations in the supra-second range is questioned.

Determinants of filled/empty optical illusion: Search for the locus of maximal effect.

A subdivided path in the visual field usually appears longer than an empty path of the same length. This phenomenon, known as the filled/empty or Oppel-Kundt illusion, depends on multiple properties of the visual stimulus, but the functional dependences have not been yet precisely characterized. We studied the illusory effect as a function of its two main determinants, the height of vertical strokes subdividing a spatial interval of a fixed length (visual angle 2.8 degree) and the number of the filling strokes, using the standard-variable distance matching paradigm. Non-monotonic dependence of the effect (over-reproduction of the spatial extension) on the varied parameters was observed in two experimental series. In the first series, the maximum effect was obtained for the fillers height roughly equal to the delimiters height (visual angle 0.25 degree); in the second series, the maximum effect was obtained for 11-13 equispaced fillers, and more accurately estimated to 15-16 as a result of a functional fit. Both data series were successfully modeled by curves generated by a single two-parametric system of form functions. Problems of determination of the maximum effect are discussed, and arguments for a genuinely multivariate approach are presented.

Characteristic changes in brain electrical activity due to chronic hypoxia in patients with obstructive sleep apnea syndrome (OSAS): a combined EEG study using LORETA and omega complexity.

EEG background activity of patients with obstructive sleep apnea syndrome (OSAS, N = 25) was compared to that of normal controls (N = 14) to reflect alterations of brain electrical activity caused by chronic intermittent hypoxia in OSAS. Global and regional (left vs. right, anterior vs. posterior) measures of spatial complexity (Omega) were used to characterize the degree of spatial synchrony of EEG. Low resolution electromagnetic tomography (LORETA) was used to localize generators of EEG activity in separate frequency bands. Comparing patients to controls, lower Omega complexity was found globally and in the right hemisphere. Using LORETA, an increased medium frequency activity was seen bilaterally in the precuneus, paracentral and posterior cingulate cortex. These findings indicate that alterations caused by chronic hypoxia in brain electrical activity in regions associated with influencing emotional regulation, long-term memory and the default mode network. Global synchronization (lower Omega complexity) may indicate a significantly reduced number of relatively independent, parallel neural processes due to chronic global hypoxic state in apneic patients as well as over the right hemisphere.

Mental states as macrostates emerging from brain electrical dynamics.

Psychophysiological correlations form the basis for different medical and scientific disciplines, but the nature of this relation has not yet been fully understood. One conceptual option is to understand the mental as "emerging" from neural processes in the specific sense that psychology and physiology provide two different descriptions of the same system. Stating these descriptions in terms of coarser- and finer-grained system states (macro- and microstates), the two descriptions may be equally adequate if the coarse-graining preserves the possibility to obtain a dynamical rule for the system. To test the empirical viability of our approach, we describe an algorithm to obtain a specific form of such a coarse-graining from data, and illustrate its operation using a simulated dynamical system. We then apply the method to an electroencephalographic recording, where we are able to identify macrostates from the physiological data that correspond to mental states of the subject.