Neutral auditory words immediately followed by painful electric shock may show reduced next-day recollection.

In this study, we investigated the effect of experimentally delivered acute pain on memory. Twenty-five participants participated in experimental sessions on consecutive days. The first session involved a categorization task to encourage memory encoding. There were two conditions, presented in randomized order, in which participants listened to a series of words, which were repeated three times. In one condition, one-third of the word items were immediately followed by a painful electrical shock. This word-shock pairing was consistent across repetition and the pain-paired items were presented unpredictably. In the other condition, all word items were not associated with pain. Response times over these repeated presentations were assessed for differences. Explicit memory was tested the following day, employing a Remember-Know assessment of word recognition, with no shocks employed. We found evidence that recollection may be reduced for pain-paired words, as the proportion of correct Remember responses (out of total correct responses) was significantly lower. There were no significant reductions in memory for non-pain items that followed painful stimulation after a period of several seconds. Consistent with the experience of pain consuming working memory resources, we theorize that painful shocks interrupt memory encoding for the immediately preceding experimental items, due to a shift in attention away from the word item.

Memory resources recover gradually over time: The effects of word frequency, presentation rate, and list composition on binding errors and mnemonic precision in source memory.

Normative word frequency has played a key role in the study of human memory, but there is little agreement as to the mechanism responsible for its effects. To determine whether word frequency affects binding probability or memory precision, we used a continuous reproduction task to examine working memory for spatial positions of words. In three experiments, after studying a list of five words, participants had to report the spatial location of one of them on a circle. Across experiments we varied word frequency, presentation rate, and the proportion of low-frequency words on each trial. A mixture model dissociated memory precision, binding failure, and guessing rate parameters from the continuous distribution of errors. On trials that contained only low- or only high-frequency words, low-frequency words led to a greater degree of error in recalling the associated location. This was due to a higher word-location binding failure and not due to differences in memory precision or guessing rates. Slowing down the presentation rate eliminated the word frequency effect by reducing binding failures for low-frequency words. Mixing frequencies in a single trial hurt high-frequency and helped low-frequency words. These findings support the idea that word frequency can lead to both positive and negative mnemonic effects depending on a trade-off between an HF encoding advantage and a LF retrieval cue advantage. We suggest that (1) low-frequency words require more resources for binding, (2) that these resources recover gradually over time, and that (3) binding fails when these resources are insufficient. (PsycInfo Database Record (c) 2022 APA, all rights reserved).

Midazolam and Ketamine Produce Distinct Neural Changes in Memory, Pain, and Fear Networks during Pain.

BACKGROUND: Despite the well-known clinical effects of midazolam and ketamine, including sedation and memory impairment, the neural mechanisms of these distinct drugs in humans are incompletely understood. The authors hypothesized that both drugs would decrease recollection memory, task-related brain activity, and long-range connectivity between components of the brain systems for memory encoding, pain processing, and fear learning. METHODS: In this randomized within-subject crossover study of 26 healthy adults, the authors used behavioral measures and functional magnetic resonance imaging to study these two anesthetics, at sedative doses, in an experimental memory paradigm using periodic pain. The primary outcome, recollection memory performance, was quantified with d' (a difference of z scores between successful recognition versus false identifications). Secondary outcomes were familiarity memory performance, serial task response times, task-related brain responses, and underlying brain connectivity from 17 preselected anatomical seed regions. All measures were determined under saline and steady-state concentrations of the drugs. RESULTS: Recollection memory was reduced under midazolam (median [95% CI], d' = 0.73 [0.43 to 1.02]) compared with saline (d' = 1.78 [1.61 to 1.96]) and ketamine (d' = 1.55 [1.12 to 1.97]; P < 0.0001). Task-related brain activity was detected under saline in areas involved in memory, pain, and fear, particularly the hippocampus, insula, and amygdala. Compared with saline, midazolam increased functional connectivity to 20 brain areas and decreased to 8, from seed regions in the precuneus, posterior cingulate, and left insula. Compared with saline, ketamine decreased connectivity to 17 brain areas and increased to 2, from 8 seed regions including the hippocampus, parahippocampus, amygdala, and anterior and primary somatosensory cortex. CONCLUSIONS: Painful stimulation during light sedation with midazolam, but not ketamine, can be accompanied by increased coherence in brain connectivity, even though details are less likely to be recollected as explicit memories.

Familiarity acts as a reduction in objective complexity.

High-complexity stimuli are thought to place extra demands on working memory when processing and manipulating such stimuli; however, operational definitions of complexity are not well established, nor are the measures that would demonstrate such effects. Here, we argue that complexity is a relative quantity that is affected by preexisting experience. Experiment 1 compared cued-recall performance for Chinese and English speakers when the stimuli involved Chinese features that varied in the number of strokes or involved Ethiopic features unfamiliar to both groups. Chinese pseudocharacters (two radicals) had half the strokes of Chinese pseudowords (two characters). The response terms were English words familiar to both groups. English speakers performed equivalently with the Ethiopic and pseudocharacters, but much worse on the pseudowords. In contrast, Chinese speakers performed equivalently with pseudowords or pseudocharacters, but worse with Ethiopic cues. Experiment 2 showed that the lack of a complexity effect for Chinese speakers was not due to greater ease of rehearsal of pseudowords compared with pseudocharacters. Experiment 3 ruled out that Chinese speakers are just better at learning paired associates involving Mandarin by demonstrating that while complexity did not affect them, other features of the stimuli did. Taken together, it appears that complexity is not an absolute property based on the number of visual elements, but rather a relative property affected by one's prior knowledge.

Greater discrimination difficulty during perceptual learning leads to stronger and more distinct representations.

Despite the conventional wisdom that it is more difficult to find a target among similar distractors, this study demonstrates that this disadvantage is short-lived, and that high target-to-distractor (TD) similarity during visual search training can have beneficial effects for learning. Participants with no prior knowledge of Chinese performed 12 hour-long sessions over 4 weeks, where they had to find a briefly presented target character among a set of distractors. At the beginning of the experiment, high TD similarity hurt performance, but the effect reversed during the first session and remained positive throughout the remaining sessions. This effect was due primarily to reducing false alarms on trials in which the target was absent from the search display. In addition, making an error on a trial with a specific character was associated with slower visual search response times on the subsequent repetition of the character, suggesting that participants paid more attention in encoding the characters after false alarms. Finally, the benefit of high TD similarity during visual search training transferred to a subsequent N-back working-memory task. These results suggest that greater discrimination difficulty likely induces stronger and more distinct representations of each character.

Frequency effects on memory: A resource-limited theory.

We present a review of frequency effects in memory, accompanied by a theory of memory, according to which the storage of new information in long-term memory (LTM) depletes a limited pool of working memory (WM) resources as an inverse function of item strength. We support the theory by showing that items with stronger representations in LTM (e.g., high frequency items) are easier to store, bind to context, and bind to one another; that WM resources are involved in storage and retrieval from LTM; that WM performance is better for stronger, more familiar stimuli. We present a novel analysis of preceding item strength, in which we show from nine existing studies that memory for an item is higher if during study it was preceded by a stronger item (e.g., a high frequency word). This effect is cumulative (the more prior items are of high frequency, the better), continuous (memory proportional to word frequency of preceding item), interacts with current item strength (larger for weaker items), and interacts with lag (decreases as the lag between the current and prior study item increases). A computational model that implements the theory is presented, which accounts for these effects. We discuss related phenomena that the model/theory can explain. (PsycINFO Database Record (c) 2019 APA, all rights reserved).

Forgetting Is a Feature, Not a Bug: Intentionally Forgetting Some Things Helps Us Remember Others by Freeing Up Working Memory Resources.

In the present study, we used an item-method directed-forgetting paradigm to test whether instructions to forget or remember one item affect memory for subsequently studied items. In two experiments (Ns = 138 and 33, respectively), recall was higher when a word pair was preceded during study by a to-be-forgotten word pair. This effect was cumulative: Performance increased when more preceding study items were to be forgotten. The effect decreased when memory was conditioned on instructions for items appearing farther back in the study list. Experiment 2 used a dual-task paradigm that suppressed, during encoding, verbal rehearsal or attentional refreshing. Neither task removed the effect, ruling out that rehearsal or attentional borrowing is responsible for the advantage conferred from previous to-be-forgotten items. We propose that memory formation depletes a limited resource that recovers over time and that to-be-forgotten items consume fewer resources, leaving more resources available for storing subsequent items. A computational model implementing the theory provided excellent fits to the data.

Memory for non-painful auditory items is influenced by whether they are experienced in a context involving painful electrical stimulation.

In this study, we sought to examine the effect of experimentally induced somatic pain on memory. Subjects heard a series of words and made categorization decisions in two different conditions. One condition included painful shocks administered just after presentation of some of the words; the other condition involved no shocks. For the condition that included painful stimulations, every other word was followed by a shock, and subjects were informed to expect this pattern. Word lists were repeated three times within each condition in randomized order, with different category judgments but consistent pain-word pairings. After a brief delay, recognition memory was assessed. Non-pain words from the pain condition were less strongly encoded than non-pain words from the completely pain-free condition. Recognition of pain-paired words was not significantly different than either subgroup of non-pain words. An important accompanying finding is that response times to repeated experimental items were slower for non-pain words from the pain condition, compared to non-pain words from the completely pain-free condition. This demonstrates that the effect of pain on memory may generalize to non-pain items experienced in the same experimental context.

The two processes underlying the testing effect- Evidence from Event-Related Potentials (ERPs).

Theoretical explanations of the testing effect (why people learn better from a test than a re-study) have largely focused on either the benefit of attempting to retrieve the answer or on the benefit of re-encoding the queried information after a successful retrieval. While a less parsimonious account, prior neuroimaging evidence has led us to postulate that both of these processes contribute to the benefit of testing over re-study. To provide further empirical support for our position, we recorded ERPs while subjects attempted to recall the second word of a pair when cued with the first. These ERPs were analyzed based on the current response accuracy and as a function of accuracy on the subsequent test, yielding three groups: the first and second tests were correct, the first was correct and the second was not, both were incorrect. Mean amplitude waveforms during the first test showed different patterns depending on the outcome patterns: Between 400 and 700 ms the amplitudes were most positive when both tests were correct and least positive when both were incorrect; mean amplitudes between 700 and 1000 ms only differed as a function of subsequent memory. They were more positive when the second test was correct. Importantly, the later component only predicted subsequent memory when the answers were not overlearned, i.e. only correctly recalled once previously. We interpret the 400-700 ms time window as a component reflecting a retrieval attempt process, which differs as a function of both current and subsequent accuracy, and the later time window as a component reflecting a re-encoding process, which only involves learning from tests, both of which are involved in the testing effect.

Item strength affects working memory capacity.

Do the processing and online manipulation of stimuli that are less familiar require more working memory (WM) resources? Is it more difficult to solve demanding problems when the symbols involved are less rather than more familiar? We explored these questions with a dual-task paradigm in which subjects had to solve algebra problems of different complexities while simultaneously holding novel symbol-digit associations in WM. The symbols were previously unknown Chinese characters, whose familiarity was manipulated by differential training frequency with a visual search task for nine hour-long sessions over 3 weeks. Subsequently, subjects solved equations that required one or two transformations. Before each trial, two different integers were assigned to two different Chinese characters of the same training frequency. Half of the time, those characters were present as variables in the equation and had to be substituted for the corresponding digits. After attempting to solve the equation, subjects had to recognize which two characters were shown immediately before that trial and to recall the integer associated with each. Solution accuracy and response times were better when the problems required one transformation only; variable substitution was not required; or the Chinese characters were high frequency. The effects of stimulus familiarity increased as the WM demands of the equation increased. Character-digit associations were also recalled less well with low-frequency characters. These results provide strong support that WM capacity depends not only on the number of chunks of information one is attempting to process but also on their strength or familiarity.

Cortical Networks Involved in Memory for Temporal Order.

We examined the neurobiological basis of temporal resetting, an aspect of temporal order memory, using a version of the delayed-match-to-multiple-sample task. While in an fMRI scanner, participants evaluated whether an item was novel or whether it had appeared before or after a reset event that signified the start of a new block of trials. Participants responded "old" to items that were repeated within the current block and "new" to both novel items and items that had last appeared before the reset event (pseudonew items). Medial-temporal, prefrontal, and occipital regions responded to absolute novelty of the stimulus-they differentiated between novel items and previously seen items, but not between old and pseudonew items. Activation for pseudonew items in the frontopolar and parietal regions, in contrast, was intermediate between old and new items. The posterior cingulate cortex extending to precuneus was the only region that showed complete temporal resetting, and its activation reflected whether an item was new or old according to the task instructions regardless of its familiarity. There was also a significant Condition (old/pseudonew) × Familiarity (second/third presentations) interaction effect on behavioral and neural measures. For pseudonew items, greater familiarity decreased response accuracy, increased RTs, increased ACC activation, and increased functional connectivity between ACC and the left frontal pole. The reverse was observed for old items. On the basis of these results, we propose a theoretical framework in which temporal resetting relies on an episodic retrieval network that is modulated by cognitive control and conflict resolution.

fMRI exploration of pedagogical benefits of repeated testing: when more is not always better.

INTRODUCTION: The testing effect refers to superior retention when study is followed by a test rather than followed by another study. Most research to date on why the testing effect occurs has been behavioral, but we employed neuroimaging methods in this study in order to shed light on the underlying processes. METHODS: Subjects were scanned while studying, restudying, and taking cued-recall tests of word pairs (with no feedback). We analyzed the BOLD responses by back sorting the encoding and test trials based on whether the subsequent test was correct or incorrect. We compared the subsequent memory patterns in initial study, restudy, and test trials. RESULTS: Overall, brain activity during test trials was a better predictor of later performance than brain activity during restudy trials. For test trials, we separately examined brain regions associated with the retrieval attempt process during successful retrieval and regions associated with the re-encoding process during retrieval in terms of prediction of subsequent memory. Regions associated with retrieval attempts were found to always predict subsequent memory success (the greater the activation, the more likely the correct recall); however, the regions associated with re-encoding would sometimes predict subsequent failure, specifically when subjects had correctly recalled the associated word several times already. CONCLUSIONS: These results suggest that whether a testing effect advantage is observed depends on both on the retrieval process and the re-encoding process which follows that retrieval.

Hitting the reset button: An ERP investigation of memory for temporal context.

This study explored how temporal context influences recognition. In an ERP experiment, subjects were asked to judge whether pictures, presented one at a time, had been seen since the previous appearance of a special reset screen. The reset screen separated sequences of successively presented stimuli and signaled a change in temporal context. A "new-repeat" picture was one that had been seen before but was to be called "new" because it had not appeared since the previous reset screen. New-repeat pictures elicited a more negative FN400 component than did "old" pictures even though both had seen before during the experiment. This suggests that familiarity, as indexed by the FN400, is sensitive to temporal context. An earlier frontopolar old/new effect distinguished pictures that were seen for the first time in the experiment from all other pictures. The late positive component (LPC), which is typically greater for old stimuli, was smaller for new-repeat pictures than for pictures seen for the first time in the experiment. Finally, individual differences in task performance were predicted by the differences in amplitude of P3b that was evoked by the onset of the reset screen.

Building knowledge requires bricks, not sand: The critical role of familiar constituents in learning.

Despite vast efforts to better understand human learning, some principles have been overlooked; specifically, that less familiar stimuli are more difficult to combine to create new knowledge and that this is because less familiar stimuli consume more working memory resources. Participants previously unfamiliar with Chinese characters were trained to discriminate visually similar characters during a visual search task over the course of a month, during which half of the characters appeared much more frequently. Ability to form associations involving these characters was tested via cued recall for novel associations consisting of two Chinese characters and an English word. Each week performance improved on the cued-recall task. Crucially, however, even though all Chinese character pairs were novel each week, those pairs consisting of more familiar characters were more easily learned. Performance on a working-memory task was better for more familiar stimuli, consistent with the claim that familiar stimuli consume fewer working memory resources. These findings have implications for optimal instruction, including second language learning.

Individual differences in working memory capacity are reflected in different ERP and EEG patterns to task difficulty.

This study examined whether there are neural markers of individual differences in working memory (WM) capacity and whether these differences are only manifest when performing a demanding WM task or at all levels of difficulty. Each subject's WM capacity was estimated using a modified digit span task prior to participation in an N-back task that varied difficulty from 1- to 4-back. While performing the N-back task, subjects wore scalp electrodes that allowed measurement of both event-related potentials (ERP) and event-related synchronization and desynchronization (ERS/ERD). Those subjects classified as low WM were more affected by the higher cognitive demands (many more errors in the 4-back task and generally slower responses) than those classified as high WM. These behavioral differences between the two groups were also apparent in the neural markers. Specifically, low WM subjects, when compared with high WM subjects, produced smaller P300 amplitudes and theta ERS, as well as greater alpha ERD at the most difficult level. Importantly, the observed differences in electrophysiological responses between the two groups were also observed at the lowest difficulty level, not just when the task challenged WM capacity. In addition, P300 amplitudes and alpha ERD responses were found to correlate with individual WM capacities independent of the task difficulty. These results suggest that there are qualitative neural differences among individuals with different WM capacities when approaching cognitive operations. Individuals with high WM capacities may make more efficient use of neural resources to keep their attention focused on the task-relevant information when performing cognitive tasks.

He who is well prepared has half won the battle: an FMRI study of task preparation.

The neural mechanism underlying preparation for tasks that vary in difficulty has not been explored. This functional magnetic resonance imaging study manipulated task difficulty by varying the working memory (WM) load of the n-back task. Each n-back task block was preceded by a preparation period involving a screen that indicated the level of difficulty of the upcoming task. Consistent with previous work, activation in some brain regions depended on WM load in the task. These regions were used as regions of interest for the univariate and multivariate (classification) analyses of preparation periods. The findings were that the patterns of brain activation during task preparation contain information about the upcoming task difficulty. (1) A support vector machine classifier was able to decode the n-back task difficulty from the patterns of brain activation during task preparation. Those individuals whose activation patterns for anticipated 1- versus 2- versus 3-back conditions were classified with higher accuracy showed better behavioral performance on the task, suggesting that task performance depends on task preparation. (2) Left inferior frontal gyrus, intraparietal sulcus, and anterior cingulate cortex parametrically decreased activation as anticipated task difficulty increased. Taken together, these results suggest dynamic involvement of the WM network not only during WM task performance, but also during task preparation.

Effective connectivity among the working memory regions during preparation for and during performance of the n-back task.

Recent neuroimaging studies have shown that working memory (WM) task difficulty can be decoded from patterns of brain activation in the WM network during preparation to perform those tasks. The inter-regional connectivity among the WM regions during task preparation has not yet been investigated. We examined this question using the graph modeling methods IMaGES and LOFS, applied to the previously published fMRI data of Manelis and Reder (2013). In that study, subjects performed 1-, 2-, and 3-back tasks. Each block of n-back was preceded by a preparation period and followed by a rest period. The analyses of task-related brain activity identified a network of 18 regions that increased in activation from 1- to 3-back (Increase network) and a network of 17 regions that decreased in activation from 1- to 3-back (Decrease network). The graph analyses revealed two types of connectivity sub-networks within the Increase and Decrease networks: "default" and "preparation-related." The "default" connectivity was present not only during task performance, but also during task preparation and during rest. We propose that this sub-network may serve as a core system that allows one to quickly activate cognitive, perceptual and motor systems in response to the relevant stimuli. The "preparation-related" connectivity was present during task preparation and task performance, but not at rest, and depended on the n-back condition. The role of this sub-network may be to pre-activate a connectivity "road map" in order to establish a top-down and bottom-up regulation of attention prior to performance on WM tasks.

Uncovering the neural mechanisms underlying learning from tests.

People learn better when re-study opportunities are replaced with tests. While researchers have begun to speculate on why testing is superior to study, few studies have directly examined the neural underpinnings of this effect. In this fMRI study, participants engaged in a study phase to learn arbitrary word pairs, followed by a cued recall test (recall second half of pair when cued with first word of pair), re-study of each pair, and finally another cycle of cued recall tests. Brain activation patterns during the first test (recall) of the studied pairs predicts performance on the second test. Importantly, while subsequent memory analyses of encoding trials also predict later accuracy, the brain regions involved in predicting later memory success are more extensive for activity during retrieval (testing) than during encoding (study). Those additional regions that predict subsequent memory based on their activation at test but not at encoding may be key to understanding the basis of the testing effect.

An attentional-adaptation account of spatial negative priming: evidence from event-related potentials.

Negative priming (NP) refers to a slower response to a target stimulus if it has been previously ignored. To examine theoretical accounts of spatial NP, we recorded behavioral measures and event-related potentials (ERPs) in a target localization task. A target and distractor briefly appeared, and the participant pressed a key corresponding to the target's location. The probability of the distractor appearing in each of four locations varied, whereas the target appeared with equal probabilities in all locations. We found that response times (RTs) were fastest when the prime distractor appeared in its most probable (frequent) location and when the prime target appeared in the location that never contained a distractor. Moreover, NP effects varied as a function of location: They were smallest when targets followed distractors in the frequent distractor location-a finding not predicted by episodic-retrieval or suppression accounts of NP. The ERP results showed that the P2, an ERP component associated with attentional orientation, was smaller in prime displays when the distractor appeared in its frequent location. Moreover, no differences were apparent between negative-prime and control trials in the N2, which is associated with suppression processes, nor in the P3, which is associated with episodic retrieval processes. These results indicate that the spatial NP effect is caused by both short- and long-term adaptation in preferences based on the history of inspecting unsuccessful locations. This article is dedicated to the memory of Edward E. Smith, and we indicate how this study was inspired by his research career.

Why it's easier to remember seeing a face we already know than one we don't: preexisting memory representations facilitate memory formation.

In two experiments, we provided support for the hypothesis that stimuli with preexisting memory representations (e.g., famous faces) are easier to associate to their encoding context than are stimuli that lack long-term memory representations (e.g., unknown faces). Subjects viewed faces superimposed on different backgrounds (e.g., the Eiffel Tower). Face recognition on a surprise memory test was better when the encoding background was reinstated than when it was swapped with a different background; however, the reinstatement advantage was modulated by how many faces had been seen with a given background, and reinstatement did not improve recognition for unknown faces. The follow-up experiment added a drug intervention that inhibited the ability to form new associations. Context reinstatement did not improve recognition for famous or unknown faces under the influence of the drug. The results suggest that it is easier to associate context to faces that have a preexisting long-term memory representation than to faces that do not.