The Neurobiology of Time Processing

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The more numerous the longer: how the integration between numerosity and time leads to a common neural response

If you are stuck in a traffic jam, the more numerous the queuing cars are, the longer you expect to wait. Time and numerosity are stimulus dimensions often associated in the same percept and whose interaction can lead to misjudgements. At brain level it is unclear to which extent time and numerosity recruit same/different neural populations and how their perceptual integration leads to changes in these populations' responses. Here we used high-spatial-resolution functional magnetic resonance imaging with neural model-based analyses to investigate how the topographic representations of numerosity and time change when these dimensions are varied together on the same visual stimulus in a congruent (the more numerous the items, the longer the display time) or incongruent manner. Compared to baseline conditions, where only one dimension was changed at a time, the variation of both stimulus dimensions led to changes in neural population responses that became more sensitive either to the two features or to one of them. Magnitude integration led also to degradation of topographies and shifts in response preferences. These changes were more pronounced in the comparison between parietal and frontal maps. Our results while pointing to partially distinct representations of time and numerosity show a common neural response to magnitude integratio

The nature of magnitude integration: Contextual interference versus active magnitude binding

Magnitude dimensions such as duration and numerosity have been shown to systematically interact, biasing each other in a congruent fashion: the more numerous a set of items is, the longer it is perceived to last in time. This integration between dimensions plays an important role in defining how we perceive magnitude. So far, however, the nature of magnitude integration remains unclear. Is magnitude integration a contextual interference, occurring whenever different types of information are concurrently available in the visual field, or does it involve an active "binding" of the different dimensions of the same object? To address these possibilities, we measured the integration bias induced by numerosity on perceived duration, in two cases: with duration and numerosity conveyed by distinct stimuli, or by the same stimulus. We show that a congruent integration effect can be observed only when the two magnitudes belong to the same stimulus. Instead, when the two magnitudes are conveyed by distinct stimuli, we observed an opposite effect. These findings demonstrate for the first time that a congruent integration occurs only between the dimensions of the same stimulus, suggesting the involvement of an active mechanism integrating the different dimensions of the same object in a unified percept

Subjective time is predicted by local and early visual processing

Time is as pervasive as it is elusive to study, and how the brain keeps track of millisecond time is still unclear. Here we addressed the mechanisms underlying duration perception by looking for a neural signature of subjective time distortion induced by motion adaptation. We recorded electroencephalographic signals in human partici-pants while they were asked to discriminate the duration of visual stimuli after different types of translational motion adaptation. Our results show that perceived duration can be predicted by the amplitude of the N200 event-related potential evoked by the adapted stimulus. Moreover, we show that the distortion of subjective time can be predicted by the activity in the Beta band frequency spectrum, at the offset of the adaptor and during the presentation of the subsequent adapted stimulus. Both effects were observed from posterior electrodes con-tralateral to the adapted stimulus. Overall, our findings suggest that local and low-level perceptual processes are involved in generating a subjective sense of time

Robust inference of causality in high-dimensional dynamical processes from the Information Imbalance of distance ranks

We introduce an approach which allows inferring causal relationships between variables for which the time evolution is available. Our method builds on the ideas of Granger Causality and Transfer Entropy, but overcomes most of their limitations. Specifically, our approach tests whether the predictability of a putative driven system Y can be improved by incorporating information from a potential driver system X, without making assumptions on the underlying dynamics and without the need to compute probability densities of the dynamic variables. Causality is assessed by a rigorous variational scheme based on the Information Imbalance of distance ranks, a recently developed statistical test capable of inferring the relative information content of different distance measures. This framework makes causality detection possible even for high-dimensional systems where only few of the variables are known or measured. Benchmark tests on coupled dynamical systems demonstrate that our approach outperforms other model-free causality detection methods, successfully handling both unidirectional and bidirectional couplings, and it is capable of detecting the arrow of time when present. We also show that the method can be used to robustly detect causality in electroencephalography data in humans.Comment: Extended acknowledgments Sectio

Human perceptual decision making of nonequilibrium fluctuations

Perceptual decision-making frequently requires making rapid, reliable choices upon encountering noisy sensory inputs. To better define the statistical processes underlying perceptual decision-making, here we characterize the choices of human participants visualizing a system of nonequilibrium stationary physical dynamics and compare such choices to the performance of an optimal agent computing Wald's sequential probability ratio test (SPRT). Participants viewed movies of a particle endowed with drifted Brownian dynamics and had to judge the motion as leftward or rightward. Overall, the results uncovered fundamental performance limits, consistent with recently established thermodynamic trade-offs involving speed, accuracy, and dissipation. Specifically, decision times are sensitive to entropy production rates. Moreover, to achieve a given level of observed accuracy, participants require more time than predicted by SPRT, indicating suboptimal integration of available information. In view of such suboptimality, we develop an alternative account based on evidence integration with a memory time constant. Setting the time constant proportionately to the deviation from equilibrium in the stimuli significantly improved trial-by-trial predictions of decision metrics with respect to SPRT. This study shows that perceptual psychophysics using stimuli rooted in nonequilibrium physical processes provides a robust platform for understanding how the brain takes decisions on stochastic information inputs.Comment: 18 pages, 13 figure

No inherent left and right side in human ‘mental number line': evidence from right brain damage

Spatial reasoning has a relevant role in mathematics and helps daily computational activities. It is widely assumed that in cultures with left-to-right reading, numbers are organized along the mental equivalent of a ruler, the mental number line, with small magnitudes located to the left of larger ones. Patients with right brain damage can disregard smaller numbers while mentally setting the midpoint of number intervals. This has been interpreted as a sign of spatial neglect for numbers on the left side of the mental number line and taken as a strong argument for the intrinsic left-to-right organization of the mental number line. Here, we put forward the understanding of this cognitive disability by discovering that patients with right brain damage disregard smaller numbers both when these are mapped on the left side of the mental number line and on the right side of an imagined clock face. This shows that the right hemisphere supports the representation of small numerical magnitudes independently from their mapping on the left or the right side of a spatial-mental layout. In addition, the study of the anatomical correlates through voxel-based lesion-symptom mapping and the mapping of lesion peaks on the diffusion tensor imaging-based reconstruction of white matter pathways showed that the rightward bias in the imagined clock-face was correlated with lesions of high-level middle temporal visual areas that code stimuli in object-centred spatial coordinates, i.e. stimuli that, like a clock face, have an inherent left and right side. In contrast, bias towards higher numbers on the mental number line was linked to white matter damage in the frontal component of the parietal-frontal number network. These anatomical findings show that the human brain does not represent the mental number line as an object with an inherent left and right side. We conclude that the bias towards higher numbers in the mental bisection of number intervals does not depend on left side spatial, imagery or object-centred neglect and that it rather depends on disruption of an abstract non-spatial representation of small numerical magnitude

Chronotopic maps in human supplementary motor area

Time is a fundamental dimension of everyday experiences. We can unmistakably sense its passage and adjust our behavior accordingly. Despite its ubiquity, the neuronal mechanisms underlying the capacity to perceive time remains unclear. Here, in two experiments using ultrahigh-field 7-Tesla (7T) functional magnetic resonance imaging (fMRI), we show that in the medial premotor cortex (supplementary motor area [SMA]) of the human brain, neural units tuned to different durations are orderly mapped in contiguous portions of the cortical surface so as to form chronomaps. The response of each portion in a chronomap is enhanced by neighboring durations and suppressed by nonpreferred durations represented in distant portions of the map. These findings suggest duration-sensitive tuning as a possible neural mechanism underlying the recognition of time and demonstrate, for the first time, that the representation of an abstract feature such as time can be instantiated by a topographical arrangement of duration-sensitive neural populations

The Role of Superior Temporal Cortex in Auditory Timing

Recently, there has been upsurge of interest in the neural mechanisms of time perception. A central question is whether the representation of time is distributed over brain regions as a function of stimulus modality, task and length of the duration used or whether it is centralized in a single specific and supramodal network. The answers seem to be converging on the former, and many areas not primarily considered as temporal processing areas remain to be investigated in the temporal domain. Here we asked whether the superior temporal gyrus, an auditory modality specific area, is involved in processing of auditory timing. Repetitive transcranial magnetic stimulation was applied over left and right superior temporal gyri while participants performed either a temporal or a frequency discrimination task of single tones. A significant decrease in performance accuracy was observed after stimulation of the right superior temporal gyrus, in addition to an increase in response uncertainty as measured by the Just Noticeable Difference. The results are specific to auditory temporal processing and performance on the frequency task was not affected. Our results further support the idea of distributed temporal processing and speak in favor of the existence of modality specific temporal regions in the human brain