Memorability shapes perceived time (and vice versa).

Visual stimuli are known to vary in their perceived duration. Some visual stimuli are also known to linger for longer in memory. Yet, whether these two features of visual processing are linked is unknown. Despite early assumptions that time is an extracted or higher-order feature of perception, more recent work over the past two decades has demonstrated that timing may be instantiated within sensory modality circuits. A primary location for many of these studies is the visual system, where duration-sensitive responses have been demonstrated. Furthermore, visual stimulus features have been observed to shift perceived duration. These findings suggest that visual circuits mediate or construct perceived time. Here we present evidence across a series of experiments that perceived time is affected by the image properties of scene size, clutter and memorability. More specifically, we observe that scene size and memorability dilate time, whereas clutter contracts it. Furthermore, the durations of more memorable images are also perceived more precisely. Conversely, the longer the perceived duration of an image, the more memorable it is. To explain these findings, we applied a recurrent convolutional neural network model of the ventral visual system, in which images are progressively processed over time. We find that more memorable images are processed faster, and that this increase in processing speed predicts both the lengthening and the increased precision of perceived durations. These findings provide evidence for a link between image features, time perception and memory that can be further explored with models of visual processing.

Separable Representations for Duration and Distance in Virtual Movements.

To navigate through the environment, humans must be able to measure both the distance traveled in space, and the interval elapsed in time. Yet, how the brain holds both of these metrics simultaneously is less well known. One possibility is that participants measure how far and how long they have traveled relative to a known reference point. To measure this, we had human participants (n = 24) perform a distance estimation task in a virtual environment in which they were cued to attend to either the spatial or temporal interval traveled while responses were measured with multiband fMRI. We observed that both dimensions evoked similar frontoparietal networks, yet with a striking rostrocaudal dissociation between temporal and spatial estimation. Multivariate classifiers trained on each dimension were further able to predict the temporal or spatial interval traveled, with centers of activation within the SMA and retrosplenial cortex for time and space, respectively. Furthermore, a cross-classification approach revealed the right supramarginal gyrus and occipital place area as regions capable of decoding the general magnitude of the traveled distance. Altogether, our findings suggest the brain uses separate systems for tracking spatial and temporal distances, which are combined together along with dimension-nonspecific estimates.

Neuroimaging Signatures of Metacognitive Improvement in Sensorimotor Timing.

Error monitoring is an essential human ability underlying learning and metacognition. In the time domain, humans possess a remarkable ability to learn and adapt to temporal intervals, yet the neural mechanisms underlying this are not clear. Recently, we demonstrated that humans improve sensorimotor time estimates when given the chance to incorporate previous trial feedback ( Bader and Wiener, 2021), suggesting that humans are metacognitively aware of their own timing errors. To test the neural basis of this metacognitive ability, human participants of both sexes underwent fMRI while they performed a visual temporal reproduction task with randomized supra-second intervals (1.5-6 s). Crucially, each trial was repeated following feedback, allowing a "re-do" to learn from the successes or errors in the initial trial. Behaviorally, we replicated our previous finding of improved re-do trial performance despite temporally uninformative (i.e., early or late) feedback. For neuroimaging, we observed a dissociation between estimating and reproducing time intervals. Estimation engaged the default mode network (DMN), including the superior frontal gyri, precuneus, and posterior cingulate, whereas reproduction activated regions associated traditionally with the "timing network" (TN), including the supplementary motor area (SMA), precentral gyrus, and right supramarginal gyrus. Notably, greater and more extensive DMN involvement was observed in re-do trials, whereas for the TN, it was more constrained. Task-based connectivity between these networks demonstrated higher inter-network correlation primarily when estimating initial trials, while re-do trial communication was higher during reproduction. Overall, these results suggest that the DMN and TN jointly mediate subjective self-awareness to improve timing performance.

Gradient Organization of Space, Time, and Numbers in the Brain: A Meta-analysis of Neuroimaging Studies.

In this study, we ran a meta-analysis of neuroimaging studies to pinpoint the neural regions that are commonly activated across space, time, and numerosity, and we tested the existence of gradient transitions among these magnitude representations in the brain. Following PRISMA guidelines, we included in the meta-analysis 112 experiments (for space domain), 114 experiments (time domain), and 115 experiments (numerosity domain), and we used the activation likelihood estimation method. We found a system of brain regions that was commonly recruited in all the three magnitudes, which included bilateral insula, the supplementary motor area (SMA), the right inferior frontal gyrus, and bilateral intraparietal sulci. Gradiental transitions between different magnitudes were found along all these regions but insulae, with space and numbers leading to gradients mainly over parietal regions (and SMA) whereas time and numbers mainly over frontal regions. These findings provide evidence for the GradiATOM theory (Gradient Theory of Magnitude), suggesting that spatial proximity given by overlapping activations and gradients is a key aspect for efficient interactions and integrations among magnitudes.

The role of consciously timed movements in shaping and improving auditory timing.

Our subjective sense of time is intertwined with a plethora of perceptual, cognitive and motor functions, and likewise, the brain is equipped to expertly filter, weight and combine these signals for seamless interactions with a dynamic world. Until relatively recently, the literature on time perception has excluded the influence of simultaneous motor activity, yet it has been found that motor circuits in the brain are at the core of most timing functions. Several studies have now identified that concurrent movements exert robust effects on perceptual timing estimates, but critically have not assessed how humans consciously judge the duration of their own movements. This creates a gap in our understanding of the mechanisms driving movement-related effects on sensory timing. We sought to address this gap by administering a sensorimotor timing task in which we explicitly compared the timing of isolated auditory tones and arm movements, or both simultaneously. We contextualized our findings within a Bayesian cue combination framework, in which separate sources of temporal information are weighted by their reliability and integrated into a unitary time estimate that is more precise than either unisensory estimate. Our results revealed differences in accuracy between auditory, movement and combined trials, and (crucially) that combined trials were the most accurately timed. Under the Bayesian framework, we found that participants' combined estimates were more precise than isolated estimates, yet were sub-optimal when compared with the model's prediction, on average. These findings elucidate previously unknown qualities of conscious motor timing and propose computational mechanisms that can describe how movements combine with perceptual signals to create unified, multimodal experiences of time.

Dissociable effects of emotional stimuli on electrophysiological indices of time and decision-making.

Previous research has demonstrated that emotional faces affect time perception, however, the underlying mechanisms are not fully understood. Earlier attempts focus on effects at the different stages of the pacemaker-accumulator model (clock, memory, and/or decision-making) including, an increase in pacemaker rate or accumulation rate via arousal or attention, respectively, or by biasing decision-making. A visual temporal bisection task with sub-second intervals was conducted in two groups to further investigate these effects; one group was strictly behavioral whereas the second included a 64-channel electroencephalogram (EEG). To separate the influence of face and timing responses, participants timed a visual stimulus, temporally flanked (before and after) by two faces, either negative or neutral, creating three trial-types: Neg→Neut, Neut→Neg, or Neut→Neut. We found a leftward shift in bisection point (BP) in Neg→Neut relative to Neut→Neut suggests an overestimation of the temporal stimulus when preceded by a negative face. Neurally, we found the face-responsive N170 was larger for negative faces and the N1 and contingent negative variation (CNV) were larger when the temporal stimulus was preceded by a negative face. Additionally, there was an interaction effect between condition and response for the late positive component of timing (LPCt) and a significant difference between response (short/long) in the neutral condition. We concluded that a preceding negative face affects the clock stage leading to more pulses being accumulated, either through attention or arousal, as indexed by a larger N1, CNV, and N170; whereas viewing a negative face after impacted decision-making mechanisms, as evidenced by the LPCt.

Selectivity of timing: A meta-analysis of temporal processing in neuroimaging studies using activation likelihood estimation and reverse inference.

Over the last few decades, many researchers have investigated time perception and how it is processed in the brain. Past studies have identified cortical and subcortical regions that play an important role in implicit and/or explicit timing tasks. In regard to timing, different regions appear to have roles of varying importance depending on the duration (sub-second vs. supra-second), type of task (such as involving motor responses or passively observing stimuli), and modality (such as auditory, visual, and sensorimotor) resulting in the literature reporting divergent results that are contingent on the specifics of the task. This meta-analysis aims at identifying regions that show activation only for explicit timing tasks through reverse inference. As such, two datasets (the first including studies that involved explicit timing tasks while the second did not) were compared using the activation likelihood estimation (ALE) algorithm. Reverse inference was implemented through Bayes factor modeling, which allowed for the comparison of the activated regions between the two ALE-maps. Results showed a constellation of regions that exhibited selective activation likelihood in explicit timing tasks with the largest posterior probability of activation resulting in the left supplementary motor area (SMA) and the bilateral insula. Some areas that have been dubbed critical for time perception in past studies (i.e., the cerebellum) did not exhibit prevalent activation after analyses.

How movements shape the perception of time.

In order to keep up with a changing environment, mobile organisms must be capable of deciding both where and when to move. This precision necessitates a strong sense of time, as otherwise we would fail in many of our movement goals. Yet, despite this intrinsic link, only recently have researchers begun to understand how these two features interact. Primarily, two effects have been observed: movements can bias time estimates, but they can also make them more precise. Here we review this literature and propose that both effects can be explained by a Bayesian cue combination framework, in which movement itself affords the most precise representation of time, which can influence perception in either feedforward or active sensing modes.

Awareness of errors and feedback in human time estimation.

Behavioral and electrophysiology studies have shown that humans possess a certain self-awareness of their individual timing ability. However, conflicting reports raise concerns about whether humans can discern the direction of their timing error, calling into question the extent of this timing awareness. To understand the depth of this ability, the impact of nondirectional feedback and reinforcement learning on time perception were examined in a unique temporal reproduction paradigm that involved a mixed set of interval durations and the opportunity to repeat every trial immediately after receiving feedback, essentially allowing a "redo." Within this task, we tested two groups of participants on versions where nondirectional feedback was provided after every response, or not provided at all. Participants in both groups demonstrated reduced central tendency and exhibited significantly greater accuracy in the redo trial temporal estimates, showcasing metacognitive ability, and an inherent capacity to adjust temporal responses despite the lack of directional information or any feedback at all. Additionally, the feedback group also exhibited an increase in the precision of responses on the redo trials, an effect not observed in the no-feedback group, suggesting that feedback may specifically reduce noise when making a temporal estimate. These findings enhance our understanding of timing self-awareness and can provide insight into what may transpire when this is disrupted.

Slowing the body slows down time perception.

Interval timing is a fundamental component of action and is susceptible to motor-related temporal distortions. Previous studies have shown that concurrent movement biases temporal estimates, but have primarily considered self-modulated movement only. However, real-world encounters often include situations in which movement is restricted or perturbed by environmental factors. In the following experiments, we introduced viscous movement environments to externally modulate movement and investigated the resulting effects on temporal perception. In two separate tasks, participants timed auditory intervals while moving a robotic arm that randomly applied four levels of viscosity. Results demonstrated that higher viscosity led to shorter perceived durations. Using a drift-diffusion model and a Bayesian observer model, we confirmed these biasing effects arose from perceptual mechanisms, instead of biases in decision making. These findings suggest that environmental perturbations are an important factor in movement-related temporal distortions, and enhance the current understanding of the interactions of motor activity and cognitive processes.

Dissociable neural indices for time and space estimates during virtual distance reproduction.

The perception and measurement of spatial and temporal dimensions have been widely studied. Yet, whether these two dimensions are processed independently is still being debated. Additionally, whether EEG components are uniquely associated with time or space, or whether they reflect a more general measure of magnitude quantity remains unknown. While undergoing EEG, subjects performed a virtual distance reproduction task, in which they were required to first walk forward for an unknown distance or time, and then reproduce that distance or time. Walking speed was varied between estimation and reproduction phases, to prevent interference between distance or time in each estimate. Behaviorally, subject performance was more variable when reproducing time than when reproducing distance, but with similar patterns of accuracy. During estimation, EEG data revealed the contingent negative variation (CNV), a measure previously associated with timing and expectation, tracked the probability of the upcoming interval, for both time and distance. However, during reproduction, the CNV exclusively oriented to the upcoming temporal interval at the start of reproduction, with no change across spatial distances. Our findings indicate that time and space are neurally separable dimensions, with the CNV both serving a supramodal role in temporal and spatial expectation, yet an exclusive role in preparing duration reproduction.

From ATOM to GradiATOM: Cortical gradients support time and space processing as revealed by a meta-analysis of neuroimaging studies.

According to the ATOM (A Theory Of Magnitude), formulated by Walsh more than fifteen years ago, there is a general system of magnitude in the brain that comprises regions, such as the parietal cortex, shared by space, time and other magnitudes. The present meta-analysis of neuroimaging studies used the Activation Likelihood Estimation (ALE) method in order to determine the set of regions commonly activated in space and time processing and to establish the neural activations specific to each magnitude domain. Following PRISMA guidelines, we included in the analysis a total of 112 and 114 experiments, exploring space and time processing, respectively. We clearly identified the presence of a system of brain regions commonly recruited in both space and time that includes: bilateral insula, the pre-supplementary motor area (pre-SMA), the right frontal operculum and the intraparietal sulci. These regions might be the best candidates to form the core magnitude neural system. Surprisingly, along each of these regions but the insula, ALE values progressed in a cortical gradient from time to space. The SMA exhibited an anterior-posterior gradient, with space activating more-anterior regions (i.e., pre-SMA) and time activating more-posterior regions (i.e., SMA-proper). Frontal and parietal regions showed a dorsal-ventral gradient: space is mediated by dorsal frontal and parietal regions, and time recruits ventral frontal and parietal regions. Our study supports but also expands the ATOM theory. Therefore, we here re-named it the 'GradiATOM' theory (Gradient Theory of Magnitude), proposing that gradient organization can facilitate the transformations and integrations of magnitude representations by allowing space- and time-related neural populations to interact with each other over minimal distances.

Modulation of Individual Alpha Frequency with tACS shifts Time Perception.

Previous studies have linked brain oscillation and timing, with evidence suggesting that alpha oscillations (10 Hz) may serve as a "sample rate" for the visual system. However, direct manipulation of alpha oscillations and time perception has not yet been demonstrated. To test this, we had 18 human subjects perform a time generalization task with visual stimuli. Additionally, we had previously recorded resting-state EEG from each subject and calculated their individual alpha frequency (IAF), estimated as the peak frequency from the mean spectrum over posterior electrodes between 8 and 13 Hz. Participants first learned a standard interval (600 ms) and were then required to judge if a new set of temporal intervals were equal or different compared with that standard. After learning the standard, participants performed this task while receiving occipital transcranial Alternating Current Stimulation (tACS). Crucially, for each subject, tACS was administered at their IAF or at off-peak alpha frequencies (IAF ± 2 Hz). Results demonstrated a linear shift in the psychometric function indicating a modification of perceived duration, such that progressively "faster" alpha stimulation led to longer perceived intervals. These results provide the first evidence that direct manipulations of alpha oscillations can shift perceived time in a manner consistent with a clock speed effect.

Movement Improves the Quality of Temporal Perception and Decision-Making.

A critical aspect of behavior is that mobile organisms must be able to precisely determine where and when to move. A better understanding of the mechanisms underlying precise movement timing and action planning is therefore crucial to understanding how we interact with the world around us. Recent evidence suggests that our experience of time is directly and intrinsically computed within the motor system, consistent with the theory of embodied cognition. To investigate the role of the motor system, we tested human subjects (n = 40) on a novel task combining reaching and time estimation. In this task, subjects were required to move a robotic manipulandum to one of two physical locations to categorize a concurrently timed suprasecond. Critically, subjects were divided into two groups: one in which movement during the interval was unrestricted and one in which they were restricted from moving until the stimulus interval had elapsed. Our results revealed a higher degree of precision for subjects in the free-moving group. A further experiment (n = 14) verified that these findings were not due to proximity to the target, counting strategies, bias, or movement length. A final experiment (n = 10) replicated these findings using a within-subjects design, performing a time reproduction task, in which movement during encoding of the interval led to more precise performance. Our findings suggest that time estimation may be instantiated within the motor system as an ongoing readout of timing judgment and confidence.

The dissociation of temporal processing behavior in concussion patients: Stable motor and dynamic perceptual timing.

Temporal processing is an integral aspect of human cognition and perception. Recent studies have suggested that patients suffering from concussion exhibit a deficit in temporal processing, characterized by poor performance on a variety of timing tasks. However, the majority of studies focusing on temporal processing deficits in concussion have focused on visual timing mechanisms. As temporal processing may be dominant for auditory-based processing, and so less susceptible to noise, we investigated patients with TBI and compared them to normal healthy controls on a battery of temporal processing tasks, including paced finger tapping and temporal bisection with sub-second intervals. The results of our investigation found that traumatic brain patients were unimpaired on the paced finger tapping task, suggesting that temporal processing deficits do not extend into motor timing and rhythmicity domain. In the temporal bisection task, TBI patients maintained precision but had a significantly higher bisection point, characterized by a greater propensity to judge stimuli as "short" and were significantly slower than controls. Analysis with a drift-diffusion model of perceptual decision-making revealed that TBI patients were specifically impaired in evidence accumulation, suggesting a smaller signal to noise ratio. Specifically, it demonstrated that patients had higher decision threshold and slower drift rates for accumulating evidence in order to arrive at a decision. Patients had to surmount higher evidence thresholds to reach a decision and were slower than controls in their rate of evidence accumulation. These results suggest specific deficits in temporal perceptual decision-making may predict the neural temporal pathways that may be compromised or unaffected, paving the way for designing targeted therapies to address these impairments.

Temporal and spatial discounting are distinct in humans.

Previous research on temporal and spatial discounting has largely focused on temporal discounting in which responses to reward stimuli are altered by the time taken to reach the reward. However, there is currently minimal research on the behavioral effects of spatial discounting. In addition, contrary to the current findings, previous research on reward discounting has suggested a correlation between temporal and spatial discounting. Here we present results from three studies, all of which employed a spatial and temporal discounting task in which subjects were immersed in a virtual reality environment and were presented with a choice between two monetary rewards, each reward varying in distance and duration. In addition, in experiments 2 and 3, the speed at which a subject could move within the virtual environment was manipulated. Our findings indicate some of the first evidence that space and time may in fact be estimated independently when discounting rewards.

A New Approach to an Old Flap: A Technique to Augment Venous Drainage from the Paramedian Forehead Flap.

The paramedian forehead flap is commonly used in nasal reconstruction, and survival of the distal part of the flap is usually essential for a good cosmetic outcome. Venous congestion leading to tissue necrosis is a recognized complication with this flap. The standard paramedian forehead flap is designed with a number of aims. These are to include the supratrochlear artery, to maximize mobility of the flap pedicle, to maximize the reach of the flap, and to minimize cosmetic implications at the donor site. The supratrochlear artery does not possess sizable venae comitantes; thus, the main pathway for venous drainage of the paramedian forehead flap is through superficial veins. The pattern and location of the superficial veins varies and therefore a standard skin pedicle design cannot be expected to always include sufficient veins to prevent venous congestion and subsequent flap necrosis. This article demonstrates the superficial venous anatomy of the forehead using computed tomographic venography, clinical demonstration, and cadaveric dissection, and describes a technique that can be carried out to augment flap venous drainage by performing careful dissection to identify additional superficial veins at the margins of the flap skin pedicle. One or more veins can then be mobilized and included with the flap pedicle to augment its venous drainage. Use of this technique should lead to a lower incidence of flap necrosis secondary to venous congestion.

An Intrinsic Role of Beta Oscillations in Memory for Time Estimation.

The neural mechanisms underlying time perception are of vital importance to a comprehensive understanding of behavior and cognition. Recent work has suggested a supramodal role for beta oscillations in measuring temporal intervals. However, the precise function of beta oscillations and whether their manipulation alters timing has yet to be determined. To accomplish this, we first re-analyzed two, separate EEG datasets and demonstrate that beta oscillations are associated with the retention and comparison of a memory standard for duration. We next conducted a study of 20 human participants using transcranial alternating current stimulation (tACS), over frontocentral cortex, at alpha and beta frequencies, during a visual temporal bisection task, finding that beta stimulation exclusively shifts the perception of time such that stimuli are reported as longer in duration. Finally, we decomposed trialwise choice data with a drift diffusion model of timing, revealing that the shift in timing is caused by a change in the starting point of accumulation, rather than the drift rate or threshold. Our results provide evidence for the intrinsic involvement of beta oscillations in the perception of time, and point to a specific role for beta oscillations in the encoding and retention of memory for temporal intervals.

A Bayesian Perspective on Accumulation in the Magnitude System.

Several theoretical and empirical work posit the existence of a common magnitude system in the brain. Such a proposal implies that manipulating stimuli in one magnitude dimension (e.g. duration in time) should interfere with the subjective estimation of another magnitude dimension (e.g. size in space). Here, we asked whether a generalized Bayesian magnitude estimation system would sample sensory evidence using a common, amodal prior. Two psychophysical experiments separately tested participants on their perception of duration, surface, and numerosity when the non-target magnitude dimensions and the rate of sensory evidence accumulation were manipulated. First, we found that duration estimation was resilient to changes in surface and numerosity, whereas lengthening (shortening) the duration yielded under- (over-) estimations of surface and numerosity. Second, the perception of surface and numerosity were affected by changes in the rate of sensory evidence accumulation, whereas duration was not. Our results suggest that a generalized magnitude system based on Bayesian computations would minimally necessitate multiple priors.

Science in the cloud (SIC): A use case in MRI connectomics.

Modern technologies are enabling scientists to collect extraordinary amounts of complex and sophisticated data across a huge range of scales like never before. With this onslaught of data, we can allow the focal point to shift from data collection to data analysis. Unfortunately, lack of standardized sharing mechanisms and practices often make reproducing or extending scientific results very difficult. With the creation of data organization structures and tools that drastically improve code portability, we now have the opportunity to design such a framework for communicating extensible scientific discoveries. Our proposed solution leverages these existing technologies and standards, and provides an accessible and extensible model for reproducible research, called 'science in the cloud' (SIC). Exploiting scientific containers, cloud computing, and cloud data services, we show the capability to compute in the cloud and run a web service that enables intimate interaction with the tools and data presented. We hope this model will inspire the community to produce reproducible and, importantly, extensible results that will enable us to collectively accelerate the rate at which scientific breakthroughs are discovered, replicated, and extended.