Neuroscientist's Behavioral Toolbox for Studying Episodic-Like Memory.

Episodic memory, the ability to recall specific events and experiences, is a cornerstone of human cognition with profound clinical implications. While animal studies have provided valuable insights into the neuronal underpinnings of episodic memory, research has largely relied on a limited subset of tasks that model only some aspects of episodic memory. In this narrative review, we provide an overview of rodent episodic-like memory tasks that expand the methodological repertoire and diversify the approaches used in episodic-like memory research. These tasks assess various aspects of human episodic memory, such as integrated what-where-when or what-where memory, source memory, free recall, temporal binding, and threshold retrieval dynamics. We review each task's general principle and consider whether alternative non-episodic mechanisms can account for the observed behavior. While our list of tasks is not exhaustive, we hope it will guide researchers in selecting models that align with their specific research objectives, leading to novel advancements and a more comprehensive understanding of mechanisms underlying specific aspects of episodic memory.

The time course of Temporal Binding in social and nonsocial interactions.

Temporal Binding (TB) is the subjective compression of action-effect intervals. While the effects of nonsocial actions are highly predictable, it is not the case when interacting with conspecifics, who often act under their own volition, at a time of their choosing. Given the relative differences in action-effect predictability in non-social and social interactions, it is plausible that TB and its properties differ across these situations. To examine this, in two experiments, we compared the time course of TB in social and nonsocial interactions, systematically varying action-effect intervals (200-2,100 ms). Participants were told they were (a) interacting with another person via a live webcam, who was in fact a confederate (social condition), (b) interacting with pre-recorded videos (nonsocial condition), or (c) observing two pre-recorded videos (control condition; Experiment 2). Results across experiments showed greater TB for social compared to nonsocial conditions, and the difference was proportional to the action-effect intervals. Further, in Experiment 1, TB was consistently observed throughout the experiment for social interactions, whereas nonsocial TB decreased from the first to the second half of the experiment. In Experiment 2, the nonsocial condition did not differ from control, whereas the social condition did, exhibiting enhanced binding. We argue these results suggest that the sociality of an interaction modulates the 'internal clock' of time perception.

Choice effects on temporal binding of action and outcomes: Examining the role of outcome focus and measures of time interval estimation.

The ability to make one's own choices is vital to the experience of intentional behavior. Such agency experiences are reflected in the perceptual compression of time between actions and resulting outcomes. Whereas some studies show that choice limitations weaken temporal binding, other studies do not find such an effect. Reviewing the literature, we noted two potential factors that may moderate choice limitation effects on temporal binding: (a) the extent to which individuals represent their actions in terms of the consequences they produce; and (b) the response mode of the time interval estimation measurement where participants report numbers or use a slider to indicate time intervals. Testing these conceptual and methodological factors in two separate experiments yielded clear effects of choice limitation on temporal binding but no clear moderator role of the two factors. Interestingly, overall analyses showed that the choice limitation effect gradually vanishes over time.

Synchrony perception across senses: A systematic review of temporal binding window changes from infancy to adolescence in typical and atypical development.

Sensory integration is increasingly acknowledged as being crucial for the development of cognitive and social abilities. However, its developmental trajectory is still little understood. This systematic review delves into the topic by investigating the literature about the developmental changes from infancy through adolescence of the Temporal Binding Window (TBW) - the epoch of time within which sensory inputs are perceived as simultaneous and therefore integrated. Following comprehensive searches across PubMed, Elsevier, and PsycInfo databases, only experimental, behavioral, English-language, peer-reviewed studies on multisensory temporal processing in 0-17-year-olds have been included. Non-behavioral, non-multisensory, and non-human studies have been excluded as those that did not directly focus on the TBW. The selection process was independently performed by two Authors. The 39 selected studies involved 2859 participants in total. Findings indicate a predisposition towards cross-modal asynchrony sensitivity and a composite, still unclear, developmental trajectory, with atypical development associated to increased asynchrony tolerance. These results highlight the need for consistent and thorough research into TBW development to inform potential interventions.

The fusion point of temporal binding: Promises and perils of multisensory accounts.

Performing an action to initiate a consequence in the environment triggers the perceptual illusion of temporal binding. This phenomenon entails that actions and following effects are perceived to occur closer in time than they do outside the action-effect relationship. Here we ask whether temporal binding can be explained in terms of multisensory integration, by assuming either multisensory fusion or partial integration of the two events. We gathered two datasets featuring a wide range of action-effect delays as a key factor influencing integration. We then tested the fit of a computational model for multisensory integration, the statistically optimal cue integration (SOCI) model. Indeed, qualitative aspects of the data on a group-level followed the principles of a multisensory account. By contrast, quantitative evidence from a comprehensive model evaluation indicated that temporal binding cannot be reduced to multisensory integration. Rather, multisensory integration should be seen as one of several component processes underlying temporal binding on an individual level.

Arousal, interindividual differences and temporal binding a psychophysiological study.

The sense of agency varies as a function of arousal in negative emotional contexts. As yet, it is unknown whether the same is true for positive affect, and how inter-individual characteristics might predict these effects. Temporal binding, an implicit measure of the sense of agency, was measured in 59 participants before and after watching either an emotionally neutral film clip or a positive film clip with high or low arousal. Analyses included participants' individual differences in subjective affective ratings, physiological arousal (pupillometry, skin conductance, heart rate), striatal dopamine levels via eye blink rates, and psychopathy. Linear mixed models showed that sexual arousal decreased temporal binding whereas calm pleasure had no facilitation effect on binding. Striatal dopamine levels were positively linked whereas subjective and physiological arousal may be negatively associated with binding towards actions. Psychopathic traits reduced the effect of high arousal on binding towards actions. These results provide evidence that individual differences influence the extent to which the temporal binding is affected by high arousing states with positive valence.

The effect of group size and task involvement on temporal binding window in clap perception.

We collect various types of information from our environment and organise it to create a coherent representation. Several researchers have suggested that multiple signals within the temporal binding window (TBW) can be integrated into a single coherent experience, such as flashes, beeps, and the McGurk effect. However, there is no evidence that TBW distortion also occurs in group interactions. This study investigates the influence of group size (i.e. the group size effect) and the degree of task involvement in temporal perception using computer-generated clap sound experiments. Participants listened to the randomly generated clap sounds and evaluated whether they were synchronised. We established three conditions based on different levels of task involvement: low (L), middle (M), and high (H) conditions. The varying task involvements reflect how participants interact with the clap sounds, ranging from passive listening in the L condition to actively generating sounds by pressing a key in the M condition, or attempting to synchronise key pressing sounds with the sounds in the H condition. Our experiments show a robust group size effect on TBW, regardless of the different conditions. In other words, as the group size increases, participants perceive the group clap as a single event. Furthermore, we found that the uncertain cause-effect relationship condition (H condition) shows the highest TBW. Interestingly, the TBW in the rigid cause-effect relationship (M condition) is the same as that in the no involvement condition (L condition). Our results suggest that a widened TBW in collective behaviour may facilitate cohesive action, enabling individuals to adapt to the group in relatively uncertain contexts.

Temporal binding and sense of agency in major depression.

Background: Alterations in the experience of controlling oneself and one's environment are of high relevance to understanding the psychopathology of depression. This study investigated the relationship between Temporal Binding for action-event sequences, sense of agency, self-efficacy and symptom severity in Major Depressive Disorder. Method: We employed the Sense of Agency Scale (SoAS) and the General Self-Efficacy Scale (GSE) to assess explicit Sense of Agency and self-efficacy in a group of 42 persons diagnosed with Major Depressive Disorder (MDD) [20 identifying as female, 19 as male; mean age 37.8 years (± 13.3)] and 40 control persons without a psychiatric diagnosis (CG) [22 identifying as female, 20 as male; mean age 38.0 years ( ± 13.3)]. Depressive symptom severity was measured using the BDI-II. We additionally performed a temporal binding paradigm as a potential correlate to Sense of Agency. Participants partook in a time estimation task judging three intervals (250ms, 450ms, 650ms) while either observing or causing stimulus presentations. The underestimation of intervals following intentional actions causing stimulus presentations (compared to merely observing the stimulus presentation) is interpreted as temporal binding. Results: SoAS scores demonstrated an inverse correlation with depressive symptoms (CG: p=.032, R2=.113; MDD: p<.001, R2=.260) and a positive correlation with GSE scores (CG: p<.001, R2=.379; MDD: p<.001, R2=.254). We found distinct differences in temporal binding between healthy participants and the Major Depressive Disorder group without significant correlation between temporal binding and the SoAS or GSE scores. The data suggest group differences in time estimation particular pertaining to time intervals involving intentional action and increasingly complex multisensory stimuli. Discussion: We investigated parameters of subjective control, namely Sense of Agency and Self Efficacy. Here, we were able to reveal their inverse relationship with depressive symptoms in patients with major depressive disorder, highlighting a profound experience of loss of control with increasing symptom load. Deficits in experiencing control, particularly involving intentional motor actions (and more complex multisensory stimuli), appear to be more pronounced in Major Depressive Disorder, involving not only negative self-efficacy expectations but also an altered Sense of Agency and temporal binding. Temporal binding and SoAS scores did not correlate, adding to the growing evidence that the two measures may not be directly related. We propose that future research be directed at this contiguous relationship between Sense of Agency and Self Efficacy in Major Depressive Disorder.

The role of self-related information in the sense of agency.

Sense of agency (SoA) refers to the subjective experience of controlling one's actions and their subsequent consequences. The present study endeavors to investigate the impact of how different degrees of self-related stimuli as action outcomes on the sense of agency by observing the temporal binding effect. Results showed that self-related sound significantly altered temporal binding, notably influencing outcome binding. A post-hoc explanation model effectively elucidated the role of self-related information in the formation of the sense of agency.

The enhancement of temporal binding effect after negative social feedback.

The present study investigated the effect of social feedback on the experiences of our actions and the outcomes (e.g. temporal binding between an action and its outcome, reflecting individuals' causal beliefs modulated by their agency judgments). In Experiment 1a, participants freely decided (voluntary action) their action timing to cause an outcome, which was followed by social feedback. A larger temporal binding (TB) following negative vs. positive events was found. This effect appeared neither in the random context where the causal belief between the action and outcome was absent (Experiment 1b) nor in the involuntary action context where participants' action timing was instructed (Experiment 1c). Experiments 2a and 2b examined the effect when the action-outcome was occluded, including reversing the order of outcome and feedback in Experiment 2b. Experiments 3a and 3b investigated the effect with only social feedback or only action-outcome presented. Results revealed that the effect found in Experiment 1 was driven by social feedback and independent of the availability of the action-outcome and the position of social feedback. Our findings demonstrate a stronger temporal integration of the action and its outcome following negative social feedback, reflecting fluctuations in sense of agency when faced with social feedback.

When Good Intention Goes Away: Social Feedback Modulates the Influence of Outcome Valence on Temporal Binding.

The retrospective view of temporal binding (TB), the temporal contraction between one's actions and their effects, proposes that TB is influenced by what happens after the action. However, the role of the interaction between multiple sources of information following the action in the formation of TB has received limited attention. The current study aims to address this gap by investigating the combined influence of social feedback and outcome valence (i.e., positive or negative outcomes) on TB. In Experiment 1, the valenced outcome was followed by either positive or negative social feedback. The results showed that negative social feedback enlarged the difference between TB over negative and positive outcomes relative to positive social feedback. The modulatory effect of social feedback on TB was not observed in the absence of causal beliefs between actions and outcomes (Experiment 2). These results highlight the importance of considering the complex interplay between cognitive, emotional, and social factors in shaping our subjective temporal experience.

Eye did this! Sense of agency in eye movements.

This study investigates the sense of agency (SoA) for saccades with implicit and explicit agency measures. In two eye tracking experiments, participants moved their eyes towards on-screen stimuli that subsequently changed color. Participants then either reproduced the temporal interval between saccade and color-change (Experiment 1) or reported the time points of these events with an auditory Libet clock (Experiment 2) to measure temporal binding effects as implicit indices of SoA. Participants were either made to believe to exert control over the color change or not (agency manipulation). Explicit ratings indicated that the manipulation of causal beliefs and hence agency was successful. However, temporal binding was only evident for caused effects, and only when a sufficiently sensitive procedure was used (auditory Libet clock). This suggests a feebler connection between temporal binding and SoA than previously proposed. The results also provide evidence for a relatively fast acquisition of sense of agency for previously never experienced types of action-effect associations. This indicates that the underlying processes of action control may be rooted in more intricate and adaptable cognitive models than previously thought. Oculomotor SoA as addressed in the present study presumably represents an important cognitive foundation of gaze-based social interaction (social sense of agency) or gaze-based human-machine interaction scenarios. PUBLIC SIGNIFICANCE STATEMENT: In this study, sense of agency for eye movements in the non-social domain is investigated in detail, using both explicit and implicit measures. Therefore, it offers novel and specific insights into comprehending sense of agency concerning effects induced by eye movements, as well as broader insights into agency pertaining to entirely newly acquired types of action-effect associations. Oculomotor sense of agency presumably represents an important cognitive foundation of gaze-based social interaction (social agency) or gaze-based human-machine interaction scenarios. Due to peculiarities of the oculomotor domain such as the varying degree of volitional control, eye movements could provide new information regarding more general theories of sense of agency in future research.

Tactile cues are more intrinsically linked to motor timing than visual cues in visual-tactile sensorimotor synchronization.

Many tasks require precise synchronization with external sensory stimuli, such as driving a car. This study investigates whether combined visual-tactile information provides additional benefits to movement synchrony over separate visual and tactile stimuli and explores the relationship with the temporal binding window for multisensory integration. In Experiment 1, participants completed a sensorimotor synchronization task to examine movement variability and a simultaneity judgment task to measure the temporal binding window. Results showed similar synchronization variability between visual-tactile and tactile-only stimuli, but significantly lower than visual only. In Experiment 2, participants completed a visual-tactile sensorimotor synchronization task with cross-modal stimuli presented inside (stimulus onset asynchrony 80 ms) and outside (stimulus-onset asynchrony 400 ms) the temporal binding window to examine temporal accuracy of movement execution. Participants synchronized their movement with the first stimulus in the cross-modal pair, either the visual or tactile stimulus. Results showed significantly greater temporal accuracy when only one stimulus was presented inside the window and the second stimulus was outside the window than when both stimuli were presented inside the window, with movement execution being more accurate when attending to the tactile stimulus. Overall, these findings indicate there may be a modality-specific benefit to sensorimotor synchronization performance, such that tactile cues are weighted more strongly than visual information as tactile information is more intrinsically linked to motor timing than visual information. Further, our findings indicate that the visual-tactile temporal binding window is related to the temporal accuracy of movement execution.

Feelings of responsibility and temporal binding: A comparison of two measures of the sense of agency.

Temporal binding refers to a subjective shortening of the interval between an action and its perceptual consequences. Temporal binding has often been used by researchers to indirectly measure participants' sense of agency (SoA), or the subjective sense of causing something to happen. Other studies have proposed links between temporal binding and feelings of moral responsibility. The present study compared subjective interval estimates to feelings of responsibility in a between-subjects design (Exp 1) and a within-subjects design (Exp 2). Participants either estimated the interval between two events (two tones in the passive condition, or a keypress followed by a tone in active conditions) or rated their feeling of responsibility for the tone(s). Manipulations of participant involvement and choice impacted feelings of responsibility more than temporal estimates. Overall, the two dependent variables followed different patterns, suggesting subjective interval estimates may not be a reliable proxy for feelings of responsibility.

Age-related differences in temporal binding and the influence of action body parts.

If voluntary action is followed by an effect with a short time delay, the time interval between action and effect is often perceived to be shorter than it actually is. This perceptual time compression is termed intentional binding or temporal binding. We investigated age-related changes in adulthood considering temporal binding and its dependence on action body parts (i.e., hand vs. foot). This experiment included 17 young adults (mean age: 21.71 ± 3.14 years) and 27 older adults (mean age: 74.41 ± 3.38 years). Participants performed a button press task using their index fingers (hand condition) or toes (foot condition). The results showed that older participants exhibited a strong time compression comparable to young participants in the voluntary condition. Older participants also showed a strong time compression in involuntary action, which was induced by a mechanical device, differently from young participants. In line with previous research, the present age-related differences in time compression considering involuntary action suggest that causal belief significantly influences event perception rather than the associated intention of action or sensory afferents. The present results also suggest that the nature of action body parts has no significant influence on temporal binding, independent of age group.

Alpha-band sensory entrainment improves audiovisual temporal acuity.

Visual and auditory stimuli are transmitted from the environment to sensory cortices with different timing, requiring the brain to encode when sensory inputs must be segregated or integrated into a single percept. The probability that different audiovisual (AV) stimuli are integrated into a single percept even when presented asynchronously is reflected in the construct of temporal binding window (TBW). There is a strong interest in testing whether it is possible to broaden or shrink TBW by using different neuromodulatory approaches that can speed up or slow down ongoing alpha oscillations, which have been repeatedly hypothesized to be an important determinant of the TBWs size. Here, we employed a web-based sensory entrainment protocol combined with a simultaneity judgment task using simple flash-beep stimuli. The aim was to test whether AV temporal acuity could be modulated trial by trial by synchronizing ongoing neural oscillations in the prestimulus period to a rhythmic sensory stream presented in the upper (∼12 Hz) or lower (∼8.5 Hz) alpha range. As a control, we implemented a nonrhythmic condition where only the first and the last entrainers were employed. Results show that upper alpha entrainment shrinks AV TBW and improves AV temporal acuity when compared with lower alpha and control conditions. Our findings represent a proof of concept of the efficacy of sensory entrainment to improve AV temporal acuity in a trial-by-trial manner, and they strengthen the idea that alpha oscillations may reflect the temporal unit of AV temporal binding.

When 2 become 1: Autistic simultaneity judgements about asynchronous audiovisual speech.

It has been proposed that autistic people experience a temporal distortion whereby the temporal binding window of multisensory integration is extended. Research to date has focused on autistic children so whether these differences persist into adulthood remains unknown. In addition, the possibility that the previous observations have arisen from between-group differences in response bias, rather than perceptual differences, has not been addressed. Participants completed simultaneity judgements of audiovisual speech stimuli across a range of stimulus-onset asynchronies. Response times and accuracy data were fitted to a drift-diffusion model so that the drift rate (a measure of processing efficiency) and starting point (response bias) could be estimated. In Experiment 1, we tested a sample of non-autistic adults who completed the Autism Quotient questionnaire. Autism Quotient score was not correlated with either drift rate or response bias, nor were there between-group differences when splitting based on the first and third quantiles of scores. In Experiment 2, we compared the performance of autistic with a group of non-autistic adults. There were no between-group differences in either drift rate or starting point. The results of this study do not support the previous suggestion that autistic people have an extended temporal binding window for audiovisual speech. In addition, exploratory analysis revealed that operationalising the temporal binding window in different ways influenced whether a group difference was observed, which is an important consideration for future work.

Age-Related Changes to Multisensory Integration and Audiovisual Speech Perception.

Multisensory integration is essential for the quick and accurate perception of our environment, particularly in everyday tasks like speech perception. Research has highlighted the importance of investigating bottom-up and top-down contributions to multisensory integration and how these change as a function of ageing. Specifically, perceptual factors like the temporal binding window and cognitive factors like attention and inhibition appear to be fundamental in the integration of visual and auditory information-integration that may become less efficient as we age. These factors have been linked to brain areas like the superior temporal sulcus, with neural oscillations in the alpha-band frequency also being implicated in multisensory processing. Age-related changes in multisensory integration may have significant consequences for the well-being of our increasingly ageing population, affecting their ability to communicate with others and safely move through their environment; it is crucial that the evidence surrounding this subject continues to be carefully investigated. This review will discuss research into age-related changes in the perceptual and cognitive mechanisms of multisensory integration and the impact that these changes have on speech perception and fall risk. The role of oscillatory alpha activity is of particular interest, as it may be key in the modulation of multisensory integration.

Age-related changes to the attentional modulation of temporal binding.

During multisensory integration, the time range within which visual and auditory information can be perceived as synchronous and bound together is known as the temporal binding window (TBW). With increasing age, the TBW becomes wider, such that older adults erroneously, and often dangerously, integrate sensory inputs that are asynchronous. Recent research suggests that attentional cues can narrow the width of the TBW in younger adults, sharpening temporal perception and increasing the accuracy of integration. However, due to their age-related declines in attentional control, it is not yet known whether older adults can deploy attentional resources to narrow the TBW in the same way as younger adults. This study investigated the age-related changes to the attentional modulation of the TBW. Thirty younger and 30 older adults completed a cued-spatial-attention version of the stream-bounce illusion, assessing the extent to which the visual and auditory stimuli were integrated when presented at three different stimulus-onset asynchronies, and when attending to a validly cued or invalidly cued location. A 2 × 2 × 3 mixed ANOVA revealed that when participants attended to the validly cued location (i.e., when attention was present), susceptibility to the stream-bounce illusion decreased. However, crucially, this attentional manipulation significantly affected audiovisual integration in younger adults, but not in older adults. These findings suggest that older adults have multisensory integration-related attentional deficits. Directions for future research and practical applications surrounding treatments to improve the safety of older adults' perception and navigation through the environment are discussed.

When did it happen? Verbal information about causal relations affects time estimation.

Usually, the closer two events occur, the more likely people infer a causal relationship between them. Recent studies have shown that this relationship between time and causality is bidirectional. Participants also tend to judge events closer in time if they assume that they are causally related. We present six experiments showing causal binding, but unlike other experiments, participants do not emit any motor action, and no physical feedback is given. Rather, all stimuli and causal information are provided verbally. After reading a list of events, participants were asked to estimate the time elapsed between two of them. Those participants who were informed that there was a causal relationship between the two events estimated them as occurring closer to each other. These results support causality- and heuristic-based explanations of temporal binding, as opposed to other explanations such as sensory integration or intentionality of action.