Lack of effects of acute exercise intensity on mnemonic discrimination.

The hippocampus is thought to support episodic memory by pattern separation, thereby supporting the ability to discriminate high similarity items. Past research evaluating whether acute exercise can improve mnemonic discrimination of high similarity items is mixed. The present experiment attempts to extend these prior mixed findings by evaluating the effects of multiple exercise intensities on hippocampal-dependent, mnemonic discrimination and memory performance. Fifty-seven young adults completed a three-condition (control, moderate-intensity, and vigorous-intensity), within-subjects crossover pretest-posttest comparison. We observed no effects of acute exercise on recognition memory or mnemonic discrimination. We discuss the implications of these null findings with the broader literature by discussing the complexity of this potential exercise-mnemonic discrimination relationship, including the unique role of exercise intensity, differences in the level of processing (e.g., conceptual vs. perceptual), and unique brain regions involved in mnemonic discrimination.

The emergence of all-or-none retrieval of chunks in verbal serial recall.

People often subdivide a list into smaller pieces, called chunks. Some theories of serial recall assume memories are stored hierarchically, with all-or-none retrieval of chunks, but most mathematical models avoid hierarchical assumptions. Johnson (Journal of Verbal Learning and Verbal Behavior, 8(6), 725-731, 1969) found steep drops in errors following correct recalls (transitional-error probabilities) within putative chunks during multi-trial letter-list learning, and viewed this as evidence for all-or-none retrieval. Here we test whether all-or-none retrieval occurs in lists studied only once. In serial recall of six-word lists (Experiment 1), transitional-error probabilities were inconsistent with all-or-none retrieval, both when participants were instructed to subdivide and when temporal grouping induced subdivision. Curiously, the same analysis of previous temporally grouped nine-letter lists produced compelling evidence for all-or-none retrieval, which may result from recoding rather than the formation of chunks. In Experiment 2, participants were pre-trained on three-word chunks. For nine-word lists constructed from those trained chunks, transitional-error probabilities exhibited more pronounced evidence of all-or-none retrieval. Nearly all effects reversed with post-cued backward recall, suggesting mechanisms that play out over the course of recall rather than encoding of the list. In sum, subdivided lists do not result in hierarchical memories after a single study trial, although they may emerge in lists formed from chunks that are previously learned as such. This suggests a continuous transition from non-hierarchical subdivision of lists to all-or-none retrieval over the course of chunk formation.

The relationship between interactive-imagery instructions and association memory.

Interactive imagery, one of the most effective strategies for remembering pairs of words, involves asking participants to form mental images during study. We tested the hypothesis that the visual image is, in fact, responsible for its memory benefit. Neither subjectively reported vividness (all experiments) nor objective imagery skill (experiments 1 and 3) could explain the benefit of interactive imagery for cued recall. Aphantasic participants, who self-identified little to no mental imagery, benefited from interactive-imagery instructions as much as controls (experiment 3). Imagery instructions did not improve memory for the constituent order of associations (AB versus BA), even when participants were told how to incorporate order within their images (experiments 1 and 2). Taken together, our results suggest that the visual format of images may not be responsible for the effectiveness of the interactive-imagery instruction and moreover, interactive imagery may not result in qualitatively different associative memories.

Competition between Associations in Memory.

If two associations share an item, one may be remembered at the expense of the other (BC recalled but not AB). Here, we identify the neural processes by which this competition materializes and is resolved. We analyzed fMRI signal while participants studied sets of pairs that reliably induced pair-to-pair associative interference, but which participants could not fully resolve. Precuneus activity tracked retrieval of previous pairs during study of later overlapping pairs. This retrieval apparently produced interference by diverting study resources from the currently displayed pair. However, when activity in ventromedial prefrontal cortex, as well as anterior subregions of the hippocampus, was present while the earlier pair had been studied, interference was reversed, and both pairs were likely to be recalled. Angular gyrus and mid-frontal activity were related to interference resolution once the participant had seen both pairs. Taken together, associations compete via precuneus-mediated competitive retrieval, but ventromedial prefrontal cortex may neutralize this by ensuring that when the earlier association is remembered while studying the later pair, memories of the two pairs can overcome interference likely via activity in mid-frontal cortex and angular gyrus.

Judgments of alphabetical order and mechanisms of congruity effects.

The congruity effect is a highly replicated feature of comparative judgments, and has been recently found in memory judgments of relative temporal order. Specifically, asking "Which came earlier?" versus "Which came later?" facilitates response times and sometimes error rates on judgments toward the beginning or end of the list, respectively. This suggests memory judgments of relative temporal order may be part of a broader class of comparative judgments. If so, the same congruity effect should also be found with the English alphabet, despite the alphabet being a longer, semantic-memory list, with forward directional encoding. A large-sample study (N = 340) produced a clear congruity effect in response time and even error rate (when controlled for response time). The large number of serial positions afforded by the alphabet enabled us to test a repertoire of mathematical models instantiating four distinct mechanisms of the congruity effect, against the empirical serial-position effects. The best-performing model assumed a response bias toward a discrete set of letters conceived of as "early" versus "late," respectively, an account that had previously been ruled out for typical comparative-judgment paradigms. In contrast, models implementing congruity effect mechanisms supported for conventional comparative judgment paradigms (based on reference-point theory or positional discriminability) produced quantitatively poorer fits, with more curvilinear serial-position effects that deviated from the data. The congruity effect thus extends to long, highly directional semantic-memory lists. However, qualitatively different serial-position effects across models suggest that, despite the superficial similarity, there are probably several quite different mechanisms that produce congruity effects, which may, in turn, depend on specific task characteristics. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Mnemonic scaffolds vary in effectiveness for serial recall.

Memory champions remember vast amounts of information in order and at first encounter by associating each study item to an anchor within a scaffold - a pre-learned, structured memory. The scaffold provides direct-access retrieval cues. Dominated by the familiar-route scaffold (Method of Loci), researchers have little insight into what characteristics of scaffolds make them effective, nor whether individual differences might play a role. We compared participant-generated mnemonic scaffolds: (a) familiar routes (Loci), (b) autobiographical stories (Story), (c) parts of the human body (Body), and (d) routine activities (Routine Activity). Loci, Body, and Story Scaffolds benefited serial recall over Control (no scaffold). The Body and Loci Scaffold were equally superior to the other scaffolds. Measures of visual imagery aptitude and vividness and body responsiveness did not predict accuracy. A second experiment tested whether embodiment could be responsible for the high level of effectiveness of the Body Scaffold; this was not supported. In short, mnemonic scaffolds are not equally effective and embodied cognition may not directly contribute to memory success. The Body Scaffold may be a strong alternative to the Method of Loci and may enhance learning for most learners, including those who do not find the Method of Loci useful.

Associative recognition without hippocampal associations.

Whereas both human and animal lesion and human neuroimaging studies have implicated the hippocampus in memory for associations, some studies find preserved associative memory following hippocampal damage. Starting with a classic summed similarity model of item recognition, we can account for associative recognition without assuming a specific hippocampally-mediated associative process. We add one key assumption: that one item can influence activation of another item's features. Feature-strength patterns, evaluated for each probe item individually, are then diagnostic of whether an item was paired with one item versus another. We suggest that feature-level inference, without explicit storage of associations, may play a critical role in associative recognition tasks. (PsycInfo Database Record (c) 2023 APA, all rights reserved).

Imagery-based strategies for memory for associations.

Cued recall of word pairs is improved by asking participants to combine items in an interactive image. Meanwhile, interactive images facilitate serial-recall (Link Method), but even better when each item is imagined alongside a previously learned peg-word (Peg List Method). We asked if a peg system could support memory for pairs, hypothesising it would outperform interactive imagery. Tested with cued recall, five study strategies were manipulated between-subjects, across two experiments: (1) Both words linked to one peg; (2) Each word linked to a different peg; (3) Peg list method but studying as a serial list; (4) Interactive imagery (within-pairs); (5) Link Method. Participants were able to apply peg-list strategies to pairs, as anticipated by mathematical modelling. Error-patterns spoke to mathematical models; peg lists exhibited distance-based confusability, characteristic of positional-coding models, and errors tended to preserve within-pair position, even for inter-item associative strategies, suggesting models of association should incorporate position. However, the peg list strategies came with a speed-accuracy tradeoff and did not challenge the superiority of the interactive imagery strategy. Without extensive practice with peg list strategies, interactive imagery remains superior for associations. Peg strategies may excel instead in tasks that primarily test serial order or with extensive training.

Emotional arousal impairs association memory: roles of prefrontal cortex regions.

The brain processes underlying impairing effects of emotional arousal on associative memory were previously attributed to two dissociable routes using high-resolution fMRI of the MTL (Madan et al. 2017). Extrahippocampal MTL regions supporting associative encoding of neutral pairs suggested unitization; conversely, associative encoding of negative pairs involved compensatory hippocampal activity. Here, whole-brain fMRI revealed prefrontal contributions: dmPFC was more involved in hippocampal-dependent negative pair learning and vmPFC in extrahippocampal neutral pair learning. Successful encoding of emotional memory associations may require emotion regulation/conflict resolution (dmPFC), while neutral memory associations may be accomplished by anchoring new information to prior knowledge (vmPFC).

The Impact of Acute Exercise Timing on Memory Interference.

This study evaluated whether the timing of acute exercise can attenuate a memory interference effect. Across two experiments, participants completed an AB/AC memory task. Participants studied eight word pairs; four denoted AB (e.g., Hero - Apple) and four control (DE) pairs. Following this List 1, participants studied eight additional word pairs (List 2); four denoted AC, re-using words from the AB pairs (e.g., Hero - Project) and four control (FG) pairs. Following their study of both lists, participants completed a cued recall assessment. In Experiment 1 (N = 100), an acute exercise bout occurred before the AB/AC memory interference task, and the participants' three lab visits (successive conditions) were control, moderate-intensity (50% HRR; heart rate reserve) exercise, and vigorous-intensity (80% HRR) exercise. In Experiment 2 (N = 68), the acute exercise occurred between List 1 and List 2, and the participants' two lab visits (successive conditions) were a (80% HRR) vigorous-intensity exercise visit and a control visit. Across both experiments, we observed evidence of both proactive and retroactive interference (p < .05), but acute exercise, regardless of intensity, did not attenuate this interference (p > .05). Acute moderate-intensity exercise was better than control or vigorous-intensity exercise in enhancing associative memory (p < .05), independent of interference. In Experiment 2, vigorous intensity exercise was associated with more pronounced interference (p < .05). Our results suggest that acute exercise can enhance associative memory performance, with no attenuation of interference by exercise.

Predicting memory from study-related brain activity.

To isolate brain activity that may reflect effective cognitive processes during the study phase of a memory task, cognitive neuroscientists commonly contrast brain activity during study of later-remembered versus later-forgotten items. This "subsequent memory effect" method has been described as identifying brain activity "predictive" of memory outcome. However, the modern field of machine learning distinguishes between descriptive analysis, subject to overfitting, and true prediction, that can classify untrained data. First, we tested whether classic event-related potential signals were, in fact, predictive of later old/new recognition memory (N = 62, 225 items/participant); this produced significant but small predictive success. Next, pattern classification of the multivariate spatiotemporal features of the single-trial EEG waveform also succeeded in predicting memory. However, the prediction was still small in magnitude. In addition, topographic maps suggested individual differences in sources of predictive activity. These findings suggest that, on average, brain activity, measured by EEG, during the study phase is only marginally predictive of subsequent memory. It is possible that this predictive approach will succeed better when other experimental factors known to influence memory outcome are also integrated into the models.NEW & NOTEWORTHY For both basic and applied reasons, an important goal is to identify brain activity present while people study materials that enable us to predict whether they will remember those materials. We show that this is possible with the conventional event-related potential "subsequent-memory-effect" signals as well as with machine learning classifiers, but only to a small degree. This is in line with behavioral research, which supports many determinants of memory apart from the cognitive processes during study.

Temporal grouping and direction of serial recall.

When lists are presented with temporal pauses between groups of items, participants' response times reiterate those pauses. Accuracy is also increased, especially at particular serial positions. By comparing forward with backward serial recall, we tested whether the influence of temporal grouping is primarily a function of serial position or output position. Results favored the latter, both when recall direction was known to participants prior to (Experiment 2) or only after (Experiment 2) studying each list. Alongside fits of variants of a temporal distinctiveness-based model, our findings suggest that the influence of temporal grouping is not just a consequence of grouping information stored during the study phase. Rather, it critically depends on participants cueing with within-chunk position during recall, combined with response suppression.

Performance on an Associative Memory Test Decreases 8 hr After Cardiovascular Exercise.

This study was designed to assess the effects of acute exercise on performance of a paired associate learning (PAL) test, an operationalization of hippocampal-dependent associative memory. Participants performed a PAL test and then ran on a treadmill (exercise group, n = 52) or solved Sudoku puzzles (control group, n = 54). Participants returned 2, 5, or 8 hr later to perform a second, different, PAL test. PAL scores for the control group did not change over time. Similarly, scores on tests taken 2 and 5 hr after exercise were not different from baseline or control data. Scores on tests taken 8 hr after exercise, however, fell significantly below baseline (by 8.6%) and control (by 9.8%) scores. These data demonstrate that acute exercise can negatively affect the encoding and retrieval of new information even hours after the exercise bout, which should be a consideration when designing exercise programs to enhance, and not hinder, learning.

Effectiveness of the method of loci is only minimally related to factors that should influence imagined navigation.

The method of loci is arguably the most famous mnemonic strategy and is highly effective for memorising lists of non-spatial information in order. As described and instructed, this strategy apparently relies on a spatial/navigational metaphor. The user imagines moving through an environment, placing (study) and reporting (recall) list items along the way. However, whether the method relies critically on this spatial/navigation metaphor is unknown. An alternative hypothesis is that the navigation component is superfluous to memory success, and the method of loci is better viewed as a special case of a larger class of imagery-based peg strategies. Training participants on three virtual environments varying in their characteristics (an apartment, an open field, and a radial-arm maze), we asked participants to use each trained environment as the basis of the method of loci to learn five 11-word lists. Performance varied significantly across environment. However, the effects were small in magnitude. Further tests suggested that navigation-relevant knowledge and ability were not major determinants of success in verbal memory, even for participants who were confirmed to have been compliant with the strategy. These findings echo neuroimaging findings that navigation-based cognition does occur during application of the method of loci, but imagined navigation is unlikely to be directly responsible for its effectiveness. Instead, the method of loci may be best viewed as a variant of peg methods.

Associative interference in older and younger adults.

Healthy older adults are more challenged by associative interference than younger adults, but prior results could have been due to differences in list discrimination ability. We used a procedure that assessed interference without requiring knowledge of list membership to test the hypothesis that older adults (60-74 years old) would show more pronounced effects of associative interference in AB/AC learning. Despite our use of a self-paced, rather than timed, study procedure, older adults performed at lower levels of accuracy than younger adults, replicating the well established associative deficit in aging (Naveh-Benjamin & Mayr, 2018). Older participants also displayed more proactive interference on average. Older participants' memory for AB and AC showed statistical independence, resembling earlier data from younger participants with a timed study procedure (Burton, Lek, & Caplan, 2017). However, younger participants, with the current self-paced procedure, produced a facilitating relationship between memory for AB and AC. Thus, younger participants not only resolved, but reversed associative interference. List discrimination could not explain these age differences. Taken together, these results extend the associative deficit in aging, finding increased susceptibility to associative proactive interference and less resolution of associative interference in older than younger participants, even when given the opportunity to compensate during self-paced study. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Reduced associative memory for negative information: impact of confidence and interactive imagery during study.

Although item-memory for emotional information is enhanced, memory for associations between items is often impaired for negative, emotionally arousing compared to neutral information. We tested two possible mechanisms underlying this impairment, using picture pairs: 1) higher confidence in one's own ability to memorise negative information may cause participants to under-study negative pairs; 2) better interactive imagery for neutral pairs could facilitate associative memory for neutral pairs more than for negative pairs. Tested with associative recognition, we replicated the impairment of associative memory for negative pairs. We also replicated the result that confidence in future memory (judgments of learning) was higher for negative than neutral pairs. Inflated confidence could not explain the impairment of associative recognition memory: Judgements of learning were positively correlated with associative memory success for both negative and neutral pairs. However, neutral pairs were rated higher in their conduciveness to interactive imagery than negative pairs, and this difference in interactive imagery showed a robust relationship to the associative memory difference. Thus, associative memory reductions for negative information are not due to differences in encoding effort. Instead, interactive imagery may be less effective for encoding of negative than neutral pairs.

A hexagonal Fourier model of grid cells.

Grid cells in rat medial entorhinal cortex are widely thought to play a major role in spatial behavior. However, the exact computational role of the population of grid cells is not known. Here we provide a descriptive model, which nonetheless considers biologically feasible mechanisms, whereby the grid cells are viewed as a two-dimensional Fourier basis set, in hexagonal coordinates, with restricted availability of basis functions. With known properties imposed in the model parameters, we demonstrate how various empirical benchmark findings are straight-forward to understand in this model. We also explain how complex computations, inherent in a Fourier model, are feasible in the medial entorhinal cortex with simple mechanisms. We further suggest, based on model experiments, that grid cells may support a form of lossy compression of contextual information, enabling its representation in an efficient manner. In sum, this hexagonal Fourier model suggests how the entire population of grid cells may be modeled in a principled way, incorporates biologically feasible mechanisms and provides a potentially powerful interpretation of the relationship between grid-cell activity and contextual information beyond spatial knowledge. This enables various phenomena to be modeled with relatively simple mechanisms, and leads to novel and testable predictions.

Emotional arousal impairs association-memory: Roles of amygdala and hippocampus.

Emotional arousal is well-known to enhance memory for individual items or events, whereas it can impair association memory. The neural mechanism of this association memory impairment by emotion is not known: In response to emotionally arousing information, amygdala activity may interfere with hippocampal associative encoding (e.g., via prefrontal cortex). Alternatively, emotional information may be harder to unitize, resulting in reduced availability of extra-hippocampal medial temporal lobe support for emotional than neutral associations. To test these opposing hypotheses, we compared neural processes underlying successful and unsuccessful encoding of emotional and neutral associations. Participants intentionally studied pairs of neutral and negative pictures (Experiments 1-3). We found reduced association-memory for negative pictures in all experiments, accompanied by item-memory increases in Experiment 2. High-resolution fMRI (Experiment 3) indicated that reductions in associative encoding of emotional information are localizable to an area in ventral-lateral amygdala, driven by attentional/salience effects in the central amygdala. Hippocampal activity was similar during both pair types, but a left hippocampal cluster related to successful encoding was observed only for negative pairs. Extra-hippocampal associative memory processes (e.g., unitization) were more effective for neutral than emotional materials. Our findings suggest that reduced emotional association memory is accompanied by increases in activity and functional coupling within the amygdala. This did not disrupt hippocampal association-memory processes, which indeed were critical for successful emotional association memory formation.

Intrusions in episodic memory: reconsolidation or interference?

It would be profoundly important if reconsolidation research in animals and other memory domains generalized to human episodic memory. A 3-d-list-discrimination procedure, based on free recall of objects, with a contextual reminder cue (the testing room), has been thought to demonstrate reconsolidation of human episodic memory (as noted in a previous study). Our goal was to replicate the central result, a high intrusion rate during recall of the target list, and evaluate the reconsolidation account relative to an alternative account, based on state-dependent learning and interference. First, replication was not straightforward (Experiment 1). Second, using a very unique, highly salient context (Experiment 2), the method produced a qualitative replication, but it was small in magnitude. A critical assumption of the reconsolidation account, that the target list is reactivated and destabilized during re-exposure to the study context, was not supported (Experiment 3). Although troubling for the reconsolidation account, the findings can be easily accommodated by an alternative account that does not assume additional neurobiological processes underlying the destabilization of consolidated memories, instead explaining intrusion rates simply in terms of well-established cognitive effects, such as item-to-context binding and interference during retrieval.

The brain's representations may be compatible with convolution-based memory models.

Convolution is a mathematical operation used in vector-models of memory that have been successful in explaining a broad range of behaviour, including memory for associations between pairs of items, an important primitive of memory upon which a broad range of everyday memory behaviour depends. However, convolution models have trouble with naturalistic item representations, which are highly auto-correlated (as one finds, e.g., with photographs), and this has cast doubt on their neural plausibility. Consequently, modellers working with convolution have used item representations composed of randomly drawn values, but introducing so-called noise-like representation raises the question how those random-like values might relate to actual item properties. We propose that a compromise solution to this problem may already exist. It has also long been known that the brain tends to reduce auto-correlations in its inputs. For example, centre-surround cells in the retina approximate a Difference-of-Gaussians (DoG) transform. This enhances edges, but also turns natural images into images that are closer to being statistically like white noise. We show the DoG-transformed images, although not optimal compared to noise-like representations, survive the convolution model better than naturalistic images. This is a proof-of-principle that the pervasive tendency of the brain to reduce auto-correlations may result in representations of information that are already adequately compatible with convolution, supporting the neural plausibility of convolution-based association-memory. (PsycINFO Database Record