Implicit visuospatial sequence representations are accessible in both the practice and the transfer hand.

A serial reaction time task was used to test whether the representations of a probabilistic second-order sequence structure are (i) stored in an effector-dependent, effector-independent intrinsic or effector-independent visuospatial code and (ii) are inter-manually accessible. Participants were trained either with the dominant or non-dominant hand. Tests were performed with both hands in the practice sequence, a random sequence, and a mirror sequence. Learning did not differ significantly between left and right-hand practice, suggesting symmetric intermanual transfer from the dominant to the non-dominant hand and vice versa. In the posttest, RTs were shorter for the practice sequence than for the random sequence, and longest for the mirror sequence. Participants were unable to freely generate or recognize the practice sequence, indicating implicit knowledge of the probabilistic sequence structure. Because sequence-specific learning did not differ significantly between hands, we conclude that representations of the probabilistic sequence structure are stored in an effector-independent visuospatial code.

The Keys to the Future? An Examination of Statistical Versus Discriminative Accounts of Serial Pattern Learning.

Sequence learning is fundamental to a wide range of cognitive functions. Explaining how sequences-and the relations between the elements they comprise-are learned is a fundamental challenge to cognitive science. However, although hundreds of articles addressing this question are published each year, the actual learning mechanisms involved in the learning of sequences are rarely investigated. We present three experiments that seek to examine these mechanisms during a typing task. Experiments 1 and 2 tested learning during typing single letters on each trial. Experiment 3 tested for "chunking" of these letters into "words." The results of these experiments were used to examine the mechanisms that could best account for them, with a focus on two particular proposals: statistical transitional probability learning and discriminative error-driven learning. Experiments 1 and 2 showed that error-driven learning was a better predictor of response latencies than either n-gram frequencies or transitional probabilities. No evidence for chunking was found in Experiment 3, probably due to interspersing visual cues with the motor response. In addition, learning occurred across a greater distance in Experiment 1 than Experiment 2, suggesting that the greater predictability that comes with increased structure leads to greater learnability. These results shed new light on the mechanism responsible for sequence learning. Despite the widely held assumption that transitional probability learning is essential to this process, the present results suggest instead that the sequences are learned through a process of discriminative learning, involving prediction and feedback from prediction error.

Development of a novel rodent rapid serial visual presentation task reveals dissociable effects of stimulant versus nonstimulant treatments on attentional processes.

The rapid serial visual presentation (RSVP) task and continuous performance tasks (CPT) are used to assess attentional impairments in patients with psychiatric and neurological conditions. This study developed a novel touchscreen task for rats based on the structure of a human RSVP task and used pharmacological manipulations to investigate their effects on different performance measures. Normal animals were trained to respond to a target image and withhold responding to distractor images presented within a continuous sequence. In a second version of the task, a false-alarm image was included, so performance could be assessed relative to two types of nontarget distractors. The effects of acute administration of stimulant and nonstimulant treatments for ADHD (amphetamine and atomoxetine) were tested in both tasks. Methylphenidate, ketamine, and nicotine were tested in the first task only. Amphetamine made animals more impulsive and decreased overall accuracy but increased accuracy when the target was presented early in the image sequence. Atomoxetine improved accuracy overall with a specific reduction in false-alarm responses and a shift in the attentional curve reflecting improved accuracy for targets later in the image sequence. However, atomoxetine also slowed responding and increased omissions. Ketamine, nicotine, and methylphenidate had no specific effects at the doses tested. These results suggest that stimulant versus nonstimulant treatments have different effects on attention and impulsive behaviour in this rat version of an RSVP task. These results also suggest that RSVP-like tasks have the potential to be used to study attention in rodents.

Serial order depends on item-dependent and item-independent contexts.

We address four issues in response to Osth and Hurlstone's (2022) commentary on the context retrieval and updating (CRU) theory of serial order (Logan, 2021). First, we clarify the relations between CRU, chains, and associations. We show that CRU is not equivalent to a chaining theory and uses similarity rather than association to retrieve contexts. Second, we fix an error Logan (2021) made in accounting for the tendency to recall ACB instead of ACD in recalling ABCDEF (fill-in vs. in-fill errors, respectively). When implemented correctly, the idea that subjects mix the current context with an initial list cue after the first order error correctly predicts that fill-in errors are more frequent than in-fill errors. Third, we address position-specific prior-list intrusions, suggesting modifications to CRU and introducing a position-coding model based on CRU representations to account for them. We suggest that position-specific prior-list intrusions are evidence for position coding on some proportion of the trials but are not evidence against item coding on other trials. Finally, we address position-specific between-group intrusions in structured lists, agreeing with Osth and Hurlstone that reasonable modifications to CRU cannot account for them. We suggest that such intrusions support position coding on some proportion of the trials but do not rule out CRU-like item-based codes. We conclude by suggesting that item-independent and item-dependent coding are alternative strategies for serial recall and we stress the importance of accounting for immediate performance. (PsycInfo Database Record (c) 2024 APA, all rights reserved).

Deconfounding serial recall: Response timing and the overarching role of grouping.

We used the timing of serial recall in several situations to reveal important aspects of recall groupings that participants construct and the reasons those groupings occur. We examined the timing of responses in the recall of digit strings within two published experiments. Cowan, Saults, Elliott, and Moreno (2002) examined memory for nine-item lists in a way that deconfounded the presentation modality, input versus output serial position (using a varied starting point of recall), memory load from items not yet recalled (using whole vs. partial recall), and the presence or absence of the temporal grouping of the lists into triads. Accuracy was strikingly different in the two modalities, with grouping drastically changing recall of acoustic lists but with little difference between grouped versus ungrouped visual lists. Despite the modality difference in recall, the present timing results show that participants imposed a similar grouped structure in recall for both modalities. When errors were made, the original grouping structure still was maintained in the output timing. Cowan et al. (2005, Experiment 1) presented ungrouped acoustic lists of varying lengths in a span task. In this case, there was never a grouping structure presented to the participant and the accuracy data did not reveal evidence of grouping, but we show clear grouping in the timing structure. This grouping structure varied with both the list length and an individual's span. Timing in both experiments shows that a grouping structure is imposed on responses in a manner that is dependent on several factors not present in the stimuli. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Exploring an online method of measuring implicit sequence-learning consciousness.

Existing methods for measuring implicit sequence-learning consciousness are conducted offline. Based on the traditional measurement of cued-generation task, this study implemented an online measurement method by converting a generation task into a forced-choice task to observe the dynamic changes of consciousness in the implicit sequence-learning process. In this study, we compared the performance of online measurement task and traditional sequence-learning tasks in 31 university students. The results revealed that the online indicators were significantly correlated with classic consciousness indicators and typical ERP components of consciousness. Without affecting the development of consciousness, the online measurement indicators were found to promptly and effectively reflect the gradually changing progression of consciousness in implicit sequence learning.

Behavioral evidence for two distinct memory systems in rats.

Serial reaction time tasks, in which subjects have to match a target to a cue, are used to explore whether non-human animals have multiple memory systems. Predictable sub-sequences embedded in the sequence of cues are responded to faster, demonstrating incidental learning, often considered implicit. Here, we used the serial implicit learning task (SILT) to determine whether rats' memory shows similar effects. In SILT, subjects must nose-poke into a sequence of two lit apertures, S1 and S2. Some S1 are always followed by the same S2, creating predictable sequences (PS). Across groups, we varied the proportion of PS trials, from 10 to 80%, and show that rats with more PS experience do better on them than on unpredictable sequences, and better than rats with less experience. We then introduced test trials in which no S2 was cued. Rats with more PS experience did better on test trials. Finally, we reversed some sequences (from predictable to unpredictable and vice versa) and changed others. We find that rats with more PS experience perseverate on old (now incorrect) responses more than those with less PS experience. Overall, we find a discontinuity in performance as the proportion of PS increases, suggesting a switch in behavioral strategies or memory systems, which we confirm using a Process Dissociation Procedure analysis. Our data suggest that rats have at least two distinct memory systems, one of which appears to be analogous to human implicit memory and is differentially activated by varying the proportion of PS in our task.

Muscle activity in explicit and implicit sequence learning: Exploring additional measures of learning and certainty via tensor decomposition.

Sequence learning in serial reaction time tasks (SRTTs) is usually inferred through the reaction time measured by a keyboard. However, this chronometric parameter offers no information beyond the time point of the button-press. We therefore examined whether sequence learning can be measured by muscle activations via electromyography (EMG) in a dual-task paradigm. The primary task was a SRTT, in which the stimuli followed a fixed sequence in some blocks, whereas the sequence was random in the control condition. The secondary task stimulus was always random. One group was informed about the fixed sequence, and the other not. We assessed three dependent variables. The chronometric parameter premotor time represents the duration between stimulus onset and the onset of EMG activity, which indicates the start of the response. The other variables describe the response itself considering the EMG activity after response start. The EMG integral was analyzed, and additionally, tensor decomposition was implemented to assess sequence dependent changes in the contribution of the obtained subcomponents. The results show explicit sequence learning in this dual-task setting. Specifically, the informed group show shorter premotor times in fixed than random sequences as well as larger EMG integral and tensor contributions. Further, increased activity seems to represent response certainty, since a decrease is found for both groups in trials following erroneous responses. Interestingly, the sensitivity to sequence and post-error effects varies between the subcomponents. The results indicate that muscle activity can be a useful indicator of response behavior in addition to chronometric parameters.

Caught in the ACTS: Defining Abstract Cognitive Task Sequences as an Independent Process.

Cognitive neuroscience currently conflates the study of serial responses (e.g., delay match to sample/nonsample, n-back) with the study of sequential operations. In this essay, our goal is to define and disentangle the latter, termed abstract cognitive task sequences (ACTS). Existing literatures address tasks requiring serial events, including procedural learning of implicit motor responses, statistical learning of predictive relationships, and judgments of attributes. These findings do not describe the behavior and underlying mechanism required to succeed at remembering to evaluate color, then shape; or to multiply, then add. A new literature is needed to characterize these sorts of second-order cognitive demands of studying a sequence of operations. Our second goal is to characterize gaps in knowledge related to ACTS that merit further investigation. In the following sections, we define more precisely what we mean by ACTS and suggest research questions that further investigation would be positioned to address.

Positional inference in rhesus macaques.

Understanding how organisms make transitive inferences is critical to understanding their general ability to learn serial relationships. In this context, transitive inference (TI) can be understood as a specific heuristic that applies broadly to many different serial learning tasks, which have been the focus of hundreds of studies involving dozens of species. In the present study, monkeys learned the order of 7-item lists of photographic stimuli by trial and error, and were then tested on "derived" lists. These derived test lists combined stimuli from multiple training lists in ambiguous ways, sometimes changing their order relative to training. We found that subjects displayed strong preferences when presented with novel test pairs, even when those pairs were drawn from different training lists. These preferences were helpful when test pairs had an ordering congruent with their ranks during training, but yielded consistently below-chance performance when pairs had an incongruent order relative to training. This behavior can be explained by the joint contributions of transitive inference and another heuristic that we refer to as "positional inference." Positional inferences play a complementary role to transitive inferences in facilitating choices between novel pairs of stimuli. The theoretical framework that best explains both transitive and positional inferences is a spatial model that represents both the position of each stimulus and its uncertainty. A computational implementation of this framework yields accurate predictions about both correct responses and errors on derived lists.

Day versus night consolidation of implicit sequence learning using manual and oculomotor activation versions of the serial reaction time task: reaction time and anticipation measures.

This study presents two experiments that explored consolidation of implicit sequence learning based on two dependent variables-reaction time (RT) and correct anticipations to clarify the role of sleep, and whether the manual component is necessary for consolidation processes. Experiment 1 (n = 37) explored the performance of adults using an ocular variant of the serial reaction time task (O-SRT) with manual activation (MA), and Experiment 2 (n = 37) used the ocular activation (OA) version of the task. Each experiment consisted of a Day and a Night group that performed two sessions of the O-SRT with an intervening 12-h offline period (morning/evening in Day group, evening/following morning in Night group). Night offline had an advantage only when manual response was required and when correct anticipations (i.e., accuracy) but not RT (i.e., speed) were measured. We associated this finding with the dual-learning processes required in the MA O-SRT that led to increased sequence specific learning overnight. When using the OA O-SRT, both groups demonstrated similar rates after offline in RT and correct anticipations. We interpreted this finding to reflect stabilization, which confirmed our hypothesis. As expected, all the groups demonstrated reduced performance when another sequence was introduced, thus reflecting sequence-specific learning. This study used a powerful procedure that allows measurement of implicit sequence learning in several ways: by evaluating two different measures (RT, correct anticipations) and by isolating different aspects of the task (i.e., with/without the manual learning component, more/less general skill learning), which are known to affect learning and consolidation.

Anterior thalamic NMDA-induced damage impairs extrapolation relying on serial stimulus patterns, in rats.

Extrapolation of serial stimulus patterns seems to depend upon identification and application of patterns relating sequences of stimuli stored in memory, thus allowing prediction of pending events never experienced before. There have been proposals that such a "generator of predictions system" would include the subiculum, mammillary bodies, anteroventral thalamus and cingulate cortex (e.g., Gray, 1982). The anteroventral thalamus (AVT) seems to be in a strategic position, both hodologically and experimentally, to allow testing of this hypothesis. This study investigated the effect of NMDA-induced damage to the anteroventral thalamus [part of the anterodorsal (AD) thalamus was also damaged in some animals], following stereotaxic minute topic microinjections, on the ability of male Wistar rats to extrapolate relying on serial stimulus patterns. Corresponding sham-operated controls received phosphate-saline buffer microinjections at the same stereotaxic coordinates. The subjects were trained to run through a straight alleyway along 31 sessions, one session per day, to get rewarded. Each session included four successive trials. Subjects exposed to the monotonic serial pattern received 14, 7, 3, 1 sunflower seeds along trials. Subjects exposed to the non-monotonic serial pattern received 14, 3, 7, 1 sunflower seeds. On the 32nd testing session, a fifth trial, never experienced before, was included immediately after the fourth trial. Sham-operated control subjects exposed to the monotonic serial pattern were expected to exhibit longer running times, since the content of their prediction in the fifth trial should be "less than 1 sunflower seeds". In contrast, control subjects exposed to the non-monotonic serial pattern were expected to exhibit shorter running times, since the content of their prediction would be "more than 1 sunflower seeds". Confirming these predictions, control subjects exposed to the monotonic serial pattern exhibited longer running times as compared to both, their own running times in previous trials within the same session and control subjects exposed to the non-monotonic schedule, thus indicating the occurrence of extrapolation. In contrast, AVT/AD lesioned subjects exposed to the monotonic schedule did not exhibit this increase in running times on the fifth trial, indicating lack of extrapolation. These results indicate that extrapolation relying on serial stimulus patterns is disrupted following extensive NMDA-induced damage to AVT and part of the AD. This represents the first consistent demonstration that the anterior thalamic nuclei are required for extrapolation of serial stimulus patterns and generation of predictions.

Encoding and consolidation of motor sequence learning in young and older adults.

Sleep benefits motor memory consolidation in young adults, but this benefit is reduced in older adults. Here we sought to understand whether differences in the neural bases of encoding between young and older adults contribute to aging-related differences in sleep-dependent consolidation of an explicit variant of the serial reaction time task (SRTT). Seventeen young and 18 older adults completed two sessions (nap, wake) one week apart. In the MRI, participants learned the SRTT. Following an afternoon interval either awake or with a nap (recorded with high-density polysomnography), performance on the SRTT was reassessed in the MRI. Imaging and behavioral results from SRTT performance showed clear sleep-dependent consolidation of motor sequence learning in older adults after a daytime nap, compared to an equal interval awake. Young adults, however, showed brain activity and behavior during encoding consistent with high SRTT performance prior to the sleep interval, and did not show further sleep-dependent performance improvements. Young adults did show reduced cortical activity following sleep, suggesting potential systems-level consolidation related to automatization. Sleep physiology data showed that sigma activity topography was affected by hippocampal and cortical activation prior to the nap in both age groups, and suggested a role of theta activity in sleep-dependent automatization in young adults. These results suggest that previously observed aging-related sleep-dependent consolidation deficits may be driven by aging-related deficiencies in fast learning processes. Here we demonstrate that when sufficient encoding strength is reached with additional training, older adults demonstrate intact sleep-dependent consolidation of motor sequence learning.

Learning to deal with delayed outcomes: EEG oscillatory and slow potentials during the prefeedback interval.

It is well established that the stimulus-preceding negativity (SPN) decreases in amplitude as a task is mastered, a phenomenon generally attributed to the reduction in anticipatory attention as feedback becomes less needed. Typically, the experiments supporting this assumption have used relatively short delays (<3 s). However, we found in a previous study that this decline in amplitude, although present during the 2.5-s prefeedback delay of a patterned key-pressing task, was absent with an 8-s delay. We reexamined this finding using a 6-s delay and found that the SPN diminished at frontal sites as participants learned a sequence of four keypress durations, but that this modulation was limited to the early half of the delay (maximum at 2 s). Decline of lateralized sensorimotor theta activity across trials was also limited to early portions of the delay. These findings suggest that processes other than anticipatory attention to feedback may be more relevant for explaining SPN diminution. Such processes could include adjustment and maintenance of action-outcome expectancies (e.g., forward models) during the prefeedback interval.

Measuring implicit sequence learning and dual task ability in mild to moderate Parkinson´s disease: A feasibility study.

We investigated the feasibility aspects of two choice reaction time tasks designed to assess implicit sequence learning and dual task ability in individuals with mild to moderate Parkinson's disease in comparison to healthy individuals. Twelve individuals with mild to moderate Parkinson's disease and 12 healthy individuals, all ≥ 60 years of age, were included. A serial reaction time task was used as a measure of implicit sequence learning and a similar task but with the addition of a simple counting task, was used as a measure of dual task ability. We have present thorough descriptive statistics of the data but we have refrained from any inferential statistics due to the small sample size. All participants understood the task instructions and the difficulty level of both tasks was deemed acceptable. There were indications of task fatigue that demand careful choices for how best to analyse the data from such tasks in future trials. Ceiling effects were present in several accuracy outcomes, but not in the reaction time outcomes. Overall, we found both tasks to be feasible to use in samples of individuals with mild to moderate Parkinson's disease and healthy older individuals.

Hippocampal and striatal responses during motor learning are modulated by prefrontal cortex stimulation.

While it is widely accepted that motor sequence learning (MSL) is supported by a prefrontal-mediated interaction between hippocampal and striatal networks, it remains unknown whether the functional responses of these networks can be modulated in humans with targeted experimental interventions. The present proof-of-concept study employed a multimodal neuroimaging approach, including functional magnetic resonance (MR) imaging and MR spectroscopy, to investigate whether individually-tailored theta-burst stimulation of the dorsolateral prefrontal cortex can modulate responses in the hippocampus and the basal ganglia during motor learning. Our results indicate that while stimulation did not modulate motor performance nor task-related brain activity, it influenced connectivity patterns within hippocampo-frontal and striatal networks. Stimulation also altered the relationship between the levels of gamma-aminobutyric acid (GABA) in the stimulated prefrontal cortex and learning-related changes in both activity and connectivity in fronto-striato-hippocampal networks. This study provides the first experimental evidence, to the best of our knowledge, that brain stimulation can alter motor learning-related functional responses in the striatum and hippocampus.

Temporally Specific Roles of Ventral Tegmental Area Projections to the Nucleus Accumbens and Prefrontal Cortex in Attention and Impulse Control.

Deficits in impulse control and attention are prominent in the symptomatology of mental disorders such as attention deficit hyperactivity disorder (ADHD), substance addiction, schizophrenia, and bipolar disorder, yet the underlying mechanisms are incompletely understood. Frontostriatal structures, such as the nucleus accumbens (NAcb), the medial prefrontal cortex (mPFC), and their dopaminergic innervation from the ventral tegmental area (VTA) have been implicated in impulse control and attention. What remains unclear is how the temporal pattern of activity of these VTA projections contributes to these processes. Here, we optogenetically stimulated VTA dopamine (DA) cells, as well as VTA projections to the NAcb core (NAcbC), NAcb shell (NAcbS), and the mPFC in male rats performing the 5-choice serial reaction time task (5-CSRTT). Our data show that stimulation of VTA DA neurons, and VTA projections to the NAcbC and the mPFC immediately before presentation of the stimulus cue, impaired attention but spared impulse control. Importantly, in addition to reducing attention, activation of VTA-NAcbS also increased impulsivity when tested under a longer intertrial interval (ITI), to provoke impulsive behavior. Optogenetic stimulation at the beginning of the ITI only partially replicated these effects. In sum, our data show how attention and impulsivity are modulated by neuronal activity in distinct ascending output pathways from the VTA in a temporally specific manner. These findings increase our understanding of the intricate mechanisms by which mesocorticolimbic circuits contribute to cognition.SIGNIFICANCE STATEMENT Deficits in impulse control and attention are prominent in the symptomatology of several mental disorders, yet the brain mechanisms involved are incompletely understood. Since frontostriatal circuits have been implicated in impulse control and attention, we here examined the role of ascending projections from the midbrain ventral tegmental area (VTA) to the nucleus accumbens (NAcb) and prefrontal cortex (PFC). Using optogenetics to individually stimulate these projections with time-locked precision, we distinguished the role that each of these projections plays, in both impulse control and attention. As such, our study enhances our understanding of the neuronal circuitry that drives impulsive and attentive behavior.

The role of the posterior parietal cortex on cognition: An exploratory study.

Theta burst stimulation (TBS) is a form of repetitive transcranial magnetic stimulation (rTMS) that can be used to increase (intermittent TBS) or reduce (continuous TBS) cortical excitability. The current study provides a preliminary report of the effects of iTBS and cTBS in healthy young adults, to investigate the causal role of the posterior parietal cortex (PPC) during the performance of four cognitive functions: attention, inhibition, sequence learning and working memory. A 2 × 2 repeated measures design was incorporated using hemisphere (left/right) and TBS type (iTBS/cTBS) as the independent variables. 20 participants performed the cognitive tasks both before and after TBS stimulation in 4 counterbalanced experimental sessions (left cTBS, right cTBS, left iTBS and right iTBS) spaced 1 week apart. No change in performance was identified for the attentional cueing task after TBS stimulation, however TBS applied to the left PPC decreased reaction time when inhibiting a reflexive response. The sequence learning task revealed differential effects for encoding of the sequence versus the learnt items. cTBS on the right hemisphere resulted in faster responses to learnt sequences, and iTBS on the right hemisphere reduced reaction times during the initial encoding of the sequence. The reaction times in the 2-back working memory task were increased when TBS stimulation was applied to the right hemisphere. Results reveal clear differential effects for tasks explored, and more specifically where TBS stimulation on right PPC could provide a potential for further investigation into improving oculomotor learning by inducing plasticity-like mechanisms in the brain.

Neurophysiological and functional neuroanatomical coding of statistical and deterministic rule information during sequence learning.

Humans are capable of acquiring multiple types of information presented in the same information stream. It has been suggested that at least two parallel learning processes are important during learning of sequential patterns-statistical learning and rule-based learning. Yet, the neurophysiological underpinnings of these parallel learning processes are not fully understood. To differentiate between the simultaneous mechanisms at the single trial level, we apply a temporal EEG signal decomposition approach together with sLORETA source localization method to delineate whether distinct statistical and rule-based learning codes can be distinguished in EEG data and can be related to distinct functional neuroanatomical structures. We demonstrate that concomitant but distinct aspects of information coded in the N2 time window play a role in these mechanisms: mismatch detection and response control underlie statistical learning and rule-based learning, respectively, albeit with different levels of time-sensitivity. Moreover, the effects of the two learning mechanisms in the different temporally decomposed clusters of neural activity also differed from each other in neural sources. Importantly, the right inferior frontal cortex (BA44) was specifically implicated in visuomotor statistical learning, confirming its role in the acquisition of transitional probabilities. In contrast, visuomotor rule-based learning was associated with the prefrontal gyrus (BA6). The results show how simultaneous learning mechanisms operate at the neurophysiological level and are orchestrated by distinct prefrontal cortical areas. The current findings deepen our understanding on the mechanisms of how humans are capable of learning multiple types of information from the same stimulus stream in a parallel fashion.

Medial prefrontal cortex and hippocampal activity differentially contribute to ordinal and temporal context retrieval during sequence memory.

Remembering sequences of events defines episodic memory, but retrieval can be driven by both ordinality and temporal contexts. Whether these modes of retrieval operate at the same time or not remains unclear. Theoretically, medial prefrontal cortex (mPFC) confers ordinality, while the hippocampus (HC) associates events in gradually changing temporal contexts. Here, we looked for evidence of each with BOLD fMRI in a sequence task that taxes both retrieval modes. To test ordinal modes, items were transferred between sequences but retained their position (e.g., AB3). Ordinal modes activated mPFC, but not HC. To test temporal contexts, we examined items that skipped ahead across lag distances (e.g., ABD). HC, but not mPFC, tracked temporal contexts. There was a mPFC and HC by retrieval mode interaction. These current results suggest that the mPFC and HC are concurrently engaged in different retrieval modes in support of remembering when an event occurred.