The timing database: An open-access, live repository for interval timing studies.

Interval timing refers to the ability to perceive and remember intervals in the seconds to minutes range. Our contemporary understanding of interval timing is derived from relatively small-scale, isolated studies that investigate a limited range of intervals with a small sample size, usually based on a single task. Consequently, the conclusions drawn from individual studies are not readily generalizable to other tasks, conditions, and task parameters. The current paper presents a live database that presents raw data from interval timing studies (currently composed of 68 datasets from eight different tasks incorporating various interval and temporal order judgments) with an online graphical user interface to easily select, compile, and download the data organized in a standard format. The Timing Database aims to promote and cultivate key and novel analyses of our timing ability by making published and future datasets accessible as open-source resources for the entire research community. In the current paper, we showcase the use of the database by testing various core ideas based on data compiled across studies (i.e., temporal accuracy, scalar property, location of the point of subjective equality, malleability of timing precision). The Timing Database will serve as the repository for interval timing studies through the submission of new datasets.

Numerical averaging in mice.

Rodents can be trained to associate different durations with different stimuli (e.g., light/sound). When the associated stimuli are presented together, maximal responding is observed around the average of individual durations (akin to averaging). The current study investigated whether mice can also average independently trained numerosities. Mice were initially trained to make 10 or 20 lever presses on a single (run) lever to obtain a reward and each fixed-ratio schedule was signaled either with an auditory or visual stimulus. Then, mice were trained to press another lever to obtain the reward after they responded on the run lever for the minimum number of presses [Fixed Consecutive Number (FCN)-10 or -20 trials] signaled by the corresponding discriminative stimulus. Following this training, FCN trials with the compound stimulus were introduced to test the counting behavior of mice when they encountered conflicting information regarding the number of responses required to obtain the reward. Our results showed that the numbers of responses on these compound test trials were around the average of the number of responses in FCN-10 and FCN-20 trials particularly when the auditory stimulus was associated with a fewer number of required responses. The counting strategy explained the behavior of the majority of the mice in the FCN-Compound test trials (as opposed to the timing strategy). The number of responses in FCN-Compound trials was accounted for equally well by the arithmetic, geometric, and Bayesian averages of the number of responses observed in FCN-10 and FCN-20 trials.

Aging impairs perceptual decision-making in mice: integrating computational and neurobiological approaches.

Decision-making is one of the cognitive domains which has been under-investigated in animal models of cognitive aging along with its neurobiological correlates. This study investigated the latent variables of the decision process using the hierarchical drift-diffusion model (HDDM). Neurobiological correlates of these processes were examined via immunohistochemistry. Young (n = 11, 4 months old), adult (n = 10, 10 months old), and old (n = 10, 18 months old) mice were tested in a perceptual decision-making task (i.e. two-alternative forced-choice; 2AFC). Observed data showed that there was an age-dependent decrease in the accuracy rate of old mice while response times were comparable between age groups. HDDM results revealed that age-dependent accuracy difference was a result of a decrease in the quality of evidence integration during decision-making. Significant positive correlations observed between evidence integration rate and the number of tyrosine hydroxylase positive (TH+) neurons in the ventral tegmental area (VTA) and axon terminals in dorsomedial striatum (DMS) suggest that decrease in the quality of evidence integration in aging is related to decreased function of mesocortical and nigrostriatal dopamine.

Interval timing deficits and their neurobiological correlates in aging mice.

Age-related neurobiological and cognitive alterations suggest that interval timing (as a related function) is also altered in aging, which can, in turn, disrupt timing-dependent functions. We investigated alterations in interval timing with aging and accompanying neurobiological changes. We tested 4-6, 10-12, and 18-20 month-old mice on the dual peak interval procedure. Results revealed a specific deficit in the termination of timed responses (stop-times). The decision processes contributed more to timing variability (vs. clock/memory process) in the aged mice. We observed age-dependent reductions in the number of dopaminergic neurons in the VTA and SNc, cholinergic neurons in the medial septum/diagonal band (MS/DB) complex, and density of dopaminergic axon terminals in the DLS/DMS. Negative correlations were found between the number of dopaminergic neurons in the VTA and stop times, and the number of cholinergic neurons in MS/DB complex and the acquisition of stop times. Our results point at age-dependent changes in the decisional components of interval timing and the role of dopaminergic and cholinergic functions in these behavioral alterations.

Monitoring line length reproduction errors.

Previous work revealed that humans can keep track of the direction and degree of errors in their temporal and numerical reproductions/estimations. Given the behavioral and psychophysical commonalities to various magnitudes and the implication of an overlapping neuroanatomical locus for their representation, we hypothesized that participants would capture the direction of errors and confidence ratings would track the magnitude of errors in line-length reproductions. In two experiments, participants reproduced various target lengths as accurately as possible, and reported the direction of their errors and provided confidence ratings for their reproductions. The isolated analysis of these two second-order judgments showed that participants can correctly report the direction of errors in their line-length reproductions and subjective confidence decreases as the magnitude of errors increases. These results show that humans can robustly keep track of the direction of errors in their line-length reproductions and their subjective confidence corroborates the magnitude of these errors.

Probabilistic Information Modulates the Timed Response Inhibition Deficit in Aging Mice.

How interval timing is affected by aging constitutes one of the contemporary research questions. There is however a limited number of studies that investigate this research question in animal models of aging. The current study investigated how temporal decision-making is affected by aging. Initially, we trained young (2-3 month-old) and old C57BL/6J male mice (18-19 month-old) independently with short (3 s) and long (9 s) intervals by signaling, in each trial, the hopper associated with the interval that is in effect in that trial. The probability of short and long trials was manipulated (0.25 or 0.75) for different animals in each age group. During testing, both hoppers were illuminated, and thus active trial type was not differentiated. We expected mice to spontaneously combine the independently acquired time interval-location-probability information to adaptively guide their timing behavior in test trials. This adaptive ability and the resultant timing behavior were analyzed and compared between the age groups. Both young and old mice indeed adjusted their timing behavior in an abrupt fashion based on the independently acquired temporal-spatial-probabilistic information. The core timing ability of old mice was also intact. However, old mice had difficulty in terminating an ongoing timed response when the probability for the short trial was higher and this difference disappeared in the group that was exposed to a lower probability of short trials. These results suggest an inhibition problem in old mice as reflected through the threshold modulation process in timed decisions, which is cognitively penetrable to the probabilistic information.

Metric error monitoring in the numerical estimates.

Recent studies have shown that participants can keep track of the magnitude and direction of their errors while reproducing target intervals (Akdogan & Balci, 2017) and producing numerosities with sequentially presented auditory stimuli (Duyan & Balci, 2018). Although the latter work demonstrated that error judgments were driven by the number rather than the total duration of sequential stimulus presentations, the number and duration of stimuli are inevitably correlated in sequential presentations. This correlation empirically limits the purity of the characterization of "numerical error monitoring". The current work expanded the scope of numerical error monitoring as a form of "metric error monitoring" to numerical estimation based on simultaneously presented array of stimuli to control for temporal correlates. Our results show that numerical error monitoring ability applies to magnitude estimation in these more controlled experimental scenarios underlining its ubiquitous nature.

Numerical error monitoring.

Error monitoring has recently been discovered to have informationally rich foundations in the timing domain. Based on the common properties of magnitude-based representations, we hypothesized that judgments on the direction and the magnitude of errors would also reflect their objective counterparts in the numerosity domain. In two experiments, we presented fast sequences of "beeps" with random interstimulus intervals and asked participants to stop the sequence when they thought the target count (7, 11, or 19) had been reached. Participants then judged how close to the target they stopped the sequence, and whether their response undershot or overshot the target. Individual linear regression fits as well as the linear mixed model with a fixed effect of reproduced numerosity on confidence ratings, and participants as independent random effects on the intercept and the slope, revealed significant positive slopes for all the target numerosities. Our results suggest that humans can keep track of the direction and degree of errors in the estimation of discrete quantities, pointing at a numerical-error-monitoring ability.

Spontaneous integration of temporal information: implications for representational/computational capacity of animals.

How do animals adapt their behaviors to changing conditions? This question relates to the debate between associative versus representational/computational approaches in cognitive science. An influential line of research that has significantly shaped the conceptual development of animal learning over decades has primarily focused on the role of associative dynamics with little-to-no ascription of representational/combinatorial capacities. The common assumption of these models is that behavioral adjustments are incremental and they result from updating of associations based on actions and their outcomes, without encoding the critical information serving as the determinant(s) of such contingencies (e.g., time in interval schedules, number in ratio schedules). On the other hand, an independent line of research provides evidence for behavioral phenomena that cannot be readily accounted for by the conventional associationist approach. In this paper, we will review different sets of findings particularly in the area of interval timing that suggest the ability of animals to make swift spontaneous computations on subjective quantities and incorporate them into their behavior. Findings of these studies constitute empirical challenges for the associationist approaches to behavioral flexibility. We argue that interval timing is a fertile ground for the formulation of critical tests of different theoretical approaches to animal behavior.