The spectro-contextual encoding and retrieval theory of episodic memory.

The spectral fingerprint hypothesis, which posits that different frequencies of oscillations underlie different cognitive operations, provides one account for how interactions between brain regions support perceptual and attentive processes (Siegel etal., 2012). Here, we explore and extend this idea to the domain of human episodic memory encoding and retrieval. Incorporating findings from the synaptic to cognitive levels of organization, we argue that spectrally precise cross-frequency coupling and phase-synchronization promote the formation of hippocampal-neocortical cell assemblies that form the basis for episodic memory. We suggest that both cell assembly firing patterns as well as the global pattern of brain oscillatory activity within hippocampal-neocortical networks represents the contents of a particular memory. Drawing upon the ideas of context reinstatement and multiple trace theory, we argue that memory retrieval is driven by internal and/or external factors which recreate these frequency-specific oscillatory patterns which occur during episodic encoding. These ideas are synthesized into a novel model of episodic memory (the spectro-contextual encoding and retrieval theory, or "SCERT") that provides several testable predictions for future research

Expected reward modulates encoding-related theta activity before an event

Oscillatory brain activity in the theta frequency range (4–8 Hz) before the onset of an event has been shown to affect the likelihood of successfully encoding the event into memory. Recent work has also indicated that frontal theta activity might be modulated by reward, but it is not clear how reward expectancy, anticipatory theta activity, and memory formation might be related. Here, we used scalp electroencephalography (EEG) to assess the relationship between these factors. EEG was recorded from healthy adults while they memorized a series of words. Each word was preceded by a cue that indicated whether a high or low monetary reward would be earned if the word was successfully remembered in a later recognition test. Frontal theta power between the presentation of the reward cue and the onset of a word was predictive of later memory for the word, but only in the high reward condition. No theta differences were observed before word onset following low reward cues. The magnitude of prestimulus encoding-related theta activity in the high reward condition was correlated with the number of high reward words that were later confidently recognized. These findings provide strong evidence for a link between reward expectancy, theta activity, and memory encoding. Theta activity before event onset seems to be especially important for the encoding of motivationally significant stimuli. One possibility is that dopaminergic activity during reward anticipation mediates frontal theta activity related to memory

Improving episodic memory: frontal-midline theta neurofeedback training increases source memory performance

Cognitive and neurofeedback training (NFT) studies have demonstrated that training-induced alterations of frontal-midline (FM) theta activity (4-8 Hz) transfer to cognitive control processes. Given that FM theta oscillations are assumed to provide top-down control for episodic memory retrieval, especially for source retrieval, that is, accurate recollection of contextual details of prior episodes, the present study investigated whether FM theta NFT transfers to memory control processes. It was assessed (1) whether FM theta NFT improves source retrieval and modulates its underlying EEG characteristics and (2) whether this transfer extends over two posttests. Over seven NFT sessions, thetraining group who trained individual FM theta activity showed greater FM theta increase than an active control group who trained randomly chosen frequency bands. The training group showed better source retrieval in a posttraining session performed 13 days after NFT and their performance increasesfrom pre- to both posttraining sessions were predicted by NFT theta increases. Thus, training-induced enhancement of memory control processes seems to protect newly formed memories from proactive interference of previously learned information. EEG analyses revealed that during pretest both groups showed source memory specific theta activity at frontal and parietal sites. Surprisingly, training-induced improvements in source retrieval tended to be accompanied by less prestimulus FM theta activity, which was predicted by NFT theta change for the training but not the control group, suggesting a more efficient use of memory control processes after training. The present findings provide unique evidence for the enhancement of memory control processes by FM theta NFT

연합기억에서의 해마의 역할: 절제 연구와 뇌파 연결성 연구로부터의 통찰

학위논문(박사) -- 서울대학교대학원 : 자연과학대학 뇌인지과학과, 2023. 8. 정천기.연합 기억은 서로 관련없는 항목들의 관계에 대한 기억으로 정의됩니다. 해마는 연합기억에서 대체할 수 없는 중요한 역할을 하는 것으로 알려져 있습니다. 그러나, 해마가 단독으로 작용하여 연합 기억을 수행하는 것은 아니라는 점에 유의하는 것은 중요합니다. 연합 기억은 뇌의 여러 영역이 상호 작용하여 작동합니다. 따라서 연합 기억을 수행할 때 단순히 특정 영역이 활성화 되는 것 보다 해마와 기억 관련 네트워크 간의 기능적 연결이 더 중요할 수 있습니다. 먼저 해마가 연합 기억에 어떤 기여를 하는지 알아보기 위해 내측 측두엽 뇌전증으로 수술을 받은 환자를 대상으로 해마의 절제 여부와 수술 후 다양한 기억력 검사에서 나타난 기억력 변화 사이의 관계를 조사했습니다. 절제 영역과 위치의 개인차를 반영하는 복셀 기반 분석을 통해 해마의 절제가 항목 기억보다는 연합 기억의 저하와 관련이 있음을 발견했습니다. 이러한 이해를 바탕으로 저는 기억의 성공과 실패를 예측하기 위해 해마와 기억 관련 대뇌 피질 네트워크 영역 사이의 단일 시행 뇌파 연결성을 활용했습니다. 그 결과, 기억의 수행도를 예측할 때 평균 90% 이상의 정확도를 달성했습니다. 이 정확도는 특정 영역의 뇌 활동만을 예측에 사용하는 것과 비교했을 때 현저히 높은 수치입니다. 요약하자면, 이 논문은 연합 기억에서 해마와 해마의 연결성의 중요한 역할을 강조합니다. 이 연구는 연합 기억 과정을 이해하는 데 있어 특정 뇌 영역에만 초점을 맞추는 것이 아니라 대규모 기억 네트워크의 역할을 이해하는 것이 중요하다는 점을 강조합니다.Associative memory refers to the ability to remember the relationships between unrelated items. The hippocampus (HC) is known to play a critical and irreplaceable role in associative memory. However, it is important to note that the HC does not operate in isolation when it comes to performing associative memory; instead, it interacts with various regions of the brain. Therefore, in the context of associative memory, the functional connectivity between the HC and memory-related networks may be more important than the mere activation of specific regions. To investigate the specific contribution of the HC to associative memory, I examined the relationship between hippocampal resection and postoperative memory changes on various memory tests in patients who underwent surgery for medial temporal lobe epilepsy (MTLE). Through a voxel-based analysis that accounts for individual differences in the resection, it was found that resection of the HC was associated with a decline in associative memory rather than item memory. This finding emphasizes the specific involvement of the HC in associative memory processes. Expanding upon this understanding, I utilized single-trial EEG connectivity between the HC and neocortical regions to predict memory success and failure. The results achieved an average accuracy of over 90% in predicting subsequent memory performance. Notably, this level of accuracy was higher compared to utilizing brain activity in specific regions. In summary, this thesis highlights the significant role of the HC and its connectivity in associative memory. It underscores the significance of hippocampal communication with large-scale brain networks, rather than solely focusing on specific brain regions, in understanding memory processes.Abstract i Contents iii List of Figures v List of Tables vi List of Abbreviations vii I. INTRODUCTION 1 1.1 Associative Memory and the Hippocampus 1 1.2 Associative Memory beyond the MTL 5 1.2.1 Successful Memory Encoding and the Default Mode Network 5 1.2.2 Subsequent Memory Effects 9 1.3 Purpose of the Present Study 13 II. METHODS 14 2.1 Participants 14 2.1.1 Experiment 1. Medial Temporal Lobe Epilepsy Patients 14 2.1.2 Experiment 2. EEG Study Participants 18 2.2 Experimental Design 19 2.2.1 Experiment 1. Pre- and Post-operative Memory Test 19 2.2.2 Expereiment 2. EEG Experimental Paradigm 20 2.3 Data Analysis 22 2.3.1 Experiment 1. MRI Image and Statistical Analysis 22 2.3.2 Experiment 2. EEG Connectivity Analysis for Memory Performance Prediction 25 III. RESULTS 30 3.1 Experiment 1. Postoperative Memory Change Analysis Results 30 3.1.2 Neuropsychological Outcome 30 3.1.3 Voxel-based Analysis 32 3.2 Experiment 2. Memory Performance Prediction Results 35 3.2.1 Behavioral Results 35 3.2.2 Differences in Connectivity Features 35 3.2.3 Classification Accuracy 35 IV. DISCUSSION 40 4.1 Summary 40 4.2 Experiment 1. Associative Memory and Hippocampal Resection 41 4.3 Experiment 2. Prediction of Associative Memory Performance Using Hippocampal Connectivity 44 4.4 Conclusion 50 V. BIBLIOGRAPHY 51 Abstract in Korean 66박

An embedded process perspective

It remains a dogma in cognitive neuroscience to separate human attention and memory into distinct modules and processes. Here we propose that brain rhythms reflect the embedded nature of these processes in the human brain, as evident from their shared neural signatures: gamma oscillations (30–90 Hz) reflect sensory information processing and activated neural representations (memory items). The theta rhythm (3–8 Hz) is a pacemaker of explicit control processes (central executive), structuring neural information processing, bit by bit, as reflected in the theta-gamma code. By representing memory items in a sequential and time-compressed manner the theta-gamma code is hypothesized to solve key problems of neural computation: (1) attentional sampling (integrating and segregating information processing), (2) mnemonic updating (implementing Hebbian learning), and (3) predictive coding (advancing information processing ahead of the real time to guide behavior). In this framework, reduced alpha oscillations (8–14 Hz) reflect activated semantic networks, involved in both explicit and implicit mnemonic processes. Linking recent theoretical accounts and empirical insights on neural rhythms to the embedded-process model advances our understanding of the integrated nature of attention and memory – as the bedrock of human cognition

The two-component model of memory development, and its potential implications for educational settings

We recently introduced a two-component model of the mechanisms underlying age differences in memory functioning across the lifespan. According to this model, memory performance is based on associative and strategic components. The associative component is relatively mature by middle childhood, whereas the strategic component shows a maturational lag and continues to develop until young adulthood. Focusing on work from our own lab, we review studies from the domains of episodic and working memory informed by this model, and discuss their potential implications for educational settings. The episodic memory studies uncover the latent potential of the associative component in childhood by documenting children's ability to greatly improve their memory performance following mnemonic instruction and training. The studies on working memory also point to an immature strategic component in children whose operation is enhanced under supportive conditions. Educational settings may aim at fostering the interplay between associative and strategic components. We explore possible routes towards this goal by linking our findings to recent trends in research on instructional design

Neural oscillations during conditional associative learning.

Associative learning requires mapping between complex stimuli and behavioural responses. When multiple stimuli are involved, conditional associative learning is a gradual process with learning based on trial and error. It is established that a distributed network of regions track associative learning, however the role of neural oscillations in human learning remains less clear. Here we used scalp EEG to test how neural oscillations change during learning of arbitrary visuo-motor associations. Participants learned to associative 48 different abstract shapes to one of four button responses through trial and error over repetitions of the shapes. To quantify how well the associations were learned for each trial, we used a state-space computational model of learning that provided a probability of each trial being correct given past performance for that stimulus, that we take as a measure of the strength of the association. We used linear modelling to relate single-trial neural oscillations to single-trial measures of association strength. We found frontal midline theta oscillations during the delay period tracked learning, where theta activity was strongest during the early stages of learning and declined as the associations were formed. Further, posterior alpha and low-beta oscillations in the cue period showed strong desynchronised activity early in learning, while stronger alpha activity during the delay period was seen as associations became well learned. Moreover, the magnitude of these effects during early learning, before the associations were learned, related to improvements in memory seen on the next presentation of the stimulus. The current study provides clear evidence that frontal theta and posterior alpha/beta oscillations play a key role during associative memory formation

Spatiotemporal Dynamics of Neural Activity During Human Episodic Memory Encoding and Retrieval

Throughout literary history, the ability to travel in time has been a source of wonder and amusement. Why this fascination with moving through time? One reason may be that people are especially attuned to the concept of time travel because we each possess our own personal mental time machine: episodic memory. Through episodic memory, we transport ourselves back in time to re-live experiences from our past. This allows us to reflect on our own self-knowledge, effectively placing ourselves in context of our lives. This dissertation investigates how our brains accomplish this highly sophisticated cognitive operation. Using a laboratory model of episodic memory (free recall) and a particularly powerful neuroimaging tool (intracranial EEG), I document the changes that occur in the brain as episodic memories are first formed and then later retrieved. I find that the episodic memory system is best conceptualized as stage-wise process consisting of distinct brain regions that activate at highly conserved times relative to memory formation/retrieval. These discrete activations are used to construct a novel neurological model of episodic memory, the Neurological Stages of Episodic Retrieval and Formation (N-SERF) model. Future work should be aimed at verifying the hypotheses put forward by the N-SERF model, we well as relating the N-SERF model to prominent computational models of episodic memory

Neural Mechanisms of Episodic Memory formation

In order to remember what you had for breakfast today, you must rely on episodic memory, the memory for personal events situated within a spatiotemporal context. In this dissertation, I use electroencephalographic (EEG) recordings to measure the neural correlates of successful episodic memory formation. The recorded EEG signals simultaneously sample local field potentials throughout the brain, and can be analyzed in terms of specific time-varying oscillatory or spectral components of neural activity which are thought to reflect the concerted activity of neuronal populations. I collected EEG recordings while participants engage in free recall, an episodic memory task during which participants must study and then recall a list of items. In the first chapter, I compare the spectral correlates during encoding of items later remembered to those later forgotten using two separate recording modalities, scalp and intracranial EEG. I find that memory formation is characterized by broad low frequency spectral power decreases and high frequency power increases across both datasets, suggesting that scalp EEG can resolve high frequency activity (HFA) and that low frequency decreases in intracranial EEG are unlikely due to pathology. In the next chapter, I connect these HFA increases to memory-specific processes by comparing study items based on how they are re- called, not whether they are recalled. I find increased HFA in left lateral cortex and hippocampus during the encoding of subsequently clustered items, those items recalled consecutively with their study neighbors at test. The precise time course of these results suggests that context updating mechanisms and item-to-context associative mechanisms support successful memory formation. In the third chapter, I measure how the formation of these episodic associations is modulated by pre-existing semantic associations by including a semantic orienting task during the encoding interval. I find that semantic processing interferes with the formation of new, episodic memories. In the final chapter, I show that the memory benefit for emotionally valenced items is better explained by a contextual mechanism than an attentional mechanism. Together, my work supports the theory that contextual encoding associative mechanisms, reflected by HFA increases in the memory network, support memory formation