Retrieval processing and episodic memory

The emergence of brain imaging has had a major impact on research into the cognitive and neural bases of human memory. An area in which this impact has been particularly strong is retrieval processing - the processes engaged when attempting to retrieve information during a memory test. Several different classes of retrieval process - such as 'mode', 'effort' and 'success' - have been invoked to account for findings from neuroimaging studies of episodic retrieval. In this article we discuss how these different kinds of process, along with a fourth kind associated with 'retrieval orientation', can be investigated in brain imaging experiments. We then review studies of retrieval processing, and assess how well their designs match up to our proposed criteria for dissociating the neural correlates of different classes of retrieval process. We conclude that few studies have used designs that permit these different kinds of process to be independently identified, and that presently there is little evidence to indicate which kinds of processing can be fractionated in terms of their neural correlates.     

Individual differences in memory span: The contribution of rehearsal, access to lexical memory,and output speed

Rehearsal speed has traditionally been seen to be the prime determinant of individual differences in memory span. Recent studies, in the main using young children as the subject population, have suggested other contributors to span performance, notably contributions from long-term memory and forgetting and retrieval processes occurring during recall. In the current research we explore individual differences in span with respect to measures of rehearsal, output time, and access to lexical memory.We replicate standard short-term phenomena; we show that the variables that influence children's span performance influence adult performance in the same way; and we show that lexical memory access appears to be a more potent source of individual differences in span than either rehearsal speed or output factors.     

Frontal-Lobe Involvement in Spatial Memory: Evidence from PET,fMRI, and Lesion Studies

Many studies have identified the prefrontal cortex as the brain area that is critical for spatial memory, both in humans and in other primates. Other studies, however, have failed to establish this relation. Therefore, the aim of this paper was to review the literature regarding the role of the human prefrontal lobe in spatial memory. This was done by examining the evidence obtained from neuropsychological patients and from studies using brain-imaging techniques (PET and fMRI). Evidence supporting the notion that the prefrontal cortex is extensively involved in spatial working memory was found. The majority of these studies, however, suggests that frontal-lobe involvement is not related to the type of material that is being processed (e.g., spatial vs. nonspatial), but to process-specific functions, such as encoding and retrieval. Theoretically, these functions could be linked to the central executive within Baddeley's working-memory model, or to recent theories that emphasize the various processes that play a role in working memory. Also, methodological issues were discussed. Further research is needed to enhance our understanding of the precise interaction of domain-specific and general processes.     

Facts, rules, and procedures in normal calculation: evidence from multiple single-patient studies of impaired arithmetic fact retrieval.

This article presents results from multiple single-case studies of brain-damaged patients with impairments in retrieval of arithmetic facts (i.e., "table" facts such as 8 x 7 = 56). The results provide a basis for exploring the types of knowledge implicated in simple arithmetic performance, the internal representations for the various knowledge types, the processes operating upon these representations, and the ways in which the representations or processes may be disrupted by brain damage.     

Metamemory Experiments in Neurological Populations: A Review

Metamemory refers to knowledge about one's memory capabilities and strategies that can aid memory, as well as the processes involved in memory self-monitoring. Although metamemory has been studied in cognitive psychology for several decades, there have been fewer studies investigating the neuropsychology of metamemory. In recent years, a growing number of studies of neurological patient groups have been conducted in order to investigate the neural correlates of metamemory. In this review, we examine the neuropsychological evidence that the frontal lobes are critically involved in monitoring and control processes, which are the central components of metamemory. The following conclusions are drawn from this literature: (1) There is a strong correlation between indices of frontal lobe function or structural integrity and metamemory accuracy (2) The combination of frontal lobe dysfunction and poor memory severely impairs metamemorial processes (3) Metamemory tasks vary in subject performance levels, and quite likely, in the underlying processes these different tasks measure, and (4) Metamemory, as measured by experimental tasks, may dissociate from basic memory retrieval processes and from global judgments of memory.     

TSUNAMI: Simultaneous Understanding,Answering, and Memory Interaction for Questions

Question processing involves parsing, memory retrieval, question categorization, initiation of appropriate answer-retrieval heuristics, answer formulation, and output. Computational and psychological models have traditionally treated these processes as separate, sequential, independent, and in pursuit of a single answer type at a time. Here this view is challenged and the implications of a theory in which question processes operate simultaneously on multiple question interpretations are explored. A highly interactive model is described in which an expectation-driven parser generates multiple question candidates, including partially-specified candidates. Question candidates act as constraints for a matcher which activates memory items. An answer retrieval process examines question candidates and the active portions of memory in an attempt to generate answer candidates. Answer candidates are examined by an output process that derives the final answer. These processes run simultaneously and interact. Three experiments on human question answering are also described which provide evidence that working memory load during question reading is affected by processes related to answer retrieval.     

Stress and memory: opposing effects of glucocorticoids on memory consolidation and memory retrieval

It is well established that glucocorticoid hormones, secreted by the adrenal cortex after a stressful event, influence cognitive performance. Some studies have found glucocorticoid-induced memory enhancement. However, many studies have reported impairing effects of glucocorticoids on memory function. This paper reviews recent findings from this laboratory on the acute effects of glucocorticoids in rats on specific memory phases, i.e., memory consolidation and memory retrieval. The evidence suggests that the consequences of glucocorticoid activation on cognition depend largely on the different memory phases investigated. Posttraining activation of glucocorticoid-sensitive pathways involving glucocorticoid receptors enhances memory consolidation in a pattern highly similar to that previously described for adrenal catecholamines. Also, similar to catecholamine effects on memory consolidation, glucocorticoid influences on memory consolidation depend on noradrenergic activation of the basolateral complex of the amygdala and interactions with other brain regions. By contrast, memory retrieval processes are usually impaired with high circulating levels of glucocorticoids or following infusions of glucocorticoid receptor agonists into the hippocampus. The hypothesis is proposed that these apparently dual effects of glucocorticoids on memory consolidation and memory retrieval might be related and that the basolateral complex of the amygdala is a key structure in a memory-modulatory system that regulates, in concert with other brain regions, stress and glucocorticoid effects on both memory consolidation and memory retrieval.     

Interleaving brain systems for episodic and recognition memory

Conflicting models persist over the nature of long-term memory. Crucial issues are whether episodic memory and recognition memory reflect the same underlying processes, and the extent to which various brain structures work as a single unit to support these processes. New findings that have resulted from improved resolution of functional brain imaging, together with recent studies of amnesia and developments in animal testing, reinforce the view that recognition memory comprises at least two independent processes: one recollective and the other using familiarity detection. Only recollective recognition appears to depend on episodic memory. Attempts to map brain areas supporting these two putative components of recognition memory indicate that they depend on separate, but interlinked, structures.     

Neural dynamics of autistic behaviors: cognitive, emotional, and timing substrates

What brain mechanisms underlie autism, and how do they give rise to autistic behavioral symptoms? This article describes a neural model, called the Imbalanced Spectrally Timed Adaptive Resonance Theory (iSTART) model, that proposes how cognitive, emotional, timing, and motor processes that involve brain regions such as the prefrontal and temporal cortex, amygdala, hippocampus, and cerebellum may interact to create and perpetuate autistic symptoms. These model processes were originally developed to explain data concerning how the brain controls normal behaviors. The iSTART model shows how autistic behavioral symptoms may arise from prescribed breakdowns in these brain processes, notably a combination of underaroused emotional depression in the amygdala and related affective brain regions, learning of hyperspecific recognition categories in the temporal and prefrontal cortices, and breakdowns of adaptively timed attentional and motor circuits in the hippocampal system and cerebellum. The model clarifies how malfunctions in a subset of these mechanisms can, through a systemwide vicious circle of environmentally mediated feedback, cause and maintain problems with them all.     

Isolating the contributions of familiarity and source information to item recognition: a time course analysis.

Recognition memory may be mediated by the retrieval of distinct types of information, notably, a general assessment of familiarity and the recovery of specific source information. A response-signal speed-accuracy trade-off variant of an exclusion procedure was used to isolate the retrieval time course for familiarity and source information. In 2 experiments, participants studied spoken and read lists (with various numbers of presentations) and then performed an exclusion task, judging an item as old only if it was in the heard list. Dual-process fits of the time course data indicated that familiarity information typically is retrieved before source information. The implications that these data have for models of recognition, including dual-process and global memory models, are discussed.     

Negative priming 1985 to 2015: a measure of inhibition,the emergence of alternative accounts,and the multiple process challenge

In this article, three generations of authors describe the background to the original article; the subsequent emergence of vigorous debates concerning what negative priming actually reflects, where radically different accounts based on memory retrieval were proposed; and a re-casting of the conceptual issues underlying studies of negative priming. What started as a simple observation (slowed reaction times) and mechanism (distractor inhibition) appears now to be best explained by a multiple mechanism account involving both episodic binding and retrieval processes as well as an inhibitory process. Emerging evidence from converging techniques such as functional magnetic resonance imaging (fMRI), and especially electroencephalography (EEG), is beginning to identify these different processes. The past 30 years of negative priming experiments has revealed the dynamic and complex cognitive processes that mediate what appear to be apparently simple behavioural effects.     

Levels of processing: a view from functional brain imaging

This paper briefly reviews two central assumptions of the levels-of-processing framework in the light of findings from recent PET and fMRI studies: First, to address the suggestion that memory traces can be seen as records of analyses carried out for the purposes of perception and comprehension, studies on encoding-retrieval overlap in brain activation patterns are considered. Second, to address the suggestion that deeper, more semantic, processing results in more durable traces, studies of how encoding activity relates to processing depth and subsequent memory performance are examined. The results show that some of the sensory regions that are activated during initial perception are subsequently reactivated during retrieval, and activity in frontal and medial-temporal brain regions is related to depth of processing and level of memory performance. Collectively, these results provide support for central components of the levels framework.     

Levels of processing: A view from functional brain imaging

This paper briefly reviews two central assumptions of the levels-of-processing framework in the light of findings from recent PET and fMRI studies: First, to address the suggestion that memory traces can be seen as records of analyses carried out for the purposes of perception and comprehension, studies on encoding-retrieval overlap in brain activation patterns are considered. Second, to address the suggestion that deeper, more semantic, processing results in more durable traces, studies of how encoding activity relates to processing depth and subsequent memory performance are examined. The results show that some of the sensory regions that are activated during initial perception are subsequently reactivated during retrieval, and activity in frontal and medial-temporal brain regions is related to depth of processing and level of memory performance. Collectively, these results provide support for central components of the levels framework.     

Neuroimaging the serial position curve. A test of single-store versus dual-store models

Neuroimaging data could help clarify the long-standing dispute between dual-store and single-store models of the serial position curve. Dual-store models assume that retrieval from late positions is dependent on short-term memory (STM), whereas retrieval from early positions is dependent on long-term memory (LTM). Single-store models, however, assume that retrieval processes for early and late items are similar, but that early items are more difficult to discriminate than late items. The present study used functional magnetic resonance imaging to examine this question. Ten young adults were scanned while they recognized items from early or late serial positions. Recognition of early items uniquely activated brain areas traditionally associated with LTM, namely, regions within the hippocampal memory system. None of these areas was activated for retrieval of late items. These results indicate differential use of LTM retrieval processes, and therefore support dual-store models over single-store models.     

Exposure to a retrieval cue in rats induces changes in regional brain glucose metabolism in the amygdala and other related brain structures

Pre-test exposure to training-related cues is known to improve subsequent retention performance. To identify brain regions engaged in processes promoted by retrieval cues, a brain imaging approach using the [6-14C]glucose autoradiographic technique was used. Sprague-Dawley rats trained in a brightness discrimination avoidance task were submitted to different cueing conditions after a 1- or a 21-day training-to-test interval (TTI). Animals were either non-cued, cued with a box, or cued with a box and the light that served as a discriminative stimulus. Effects of the different cueing conditions on retention performance or on metabolic activity in 58 different brain regions were investigated. Rats cued with the light exhibited a subsequent improvement of their retention performance relative to controls, when tested at the 1- but not 21-days TTI, confirming our previous results. At the 1-day retention interval, a comparison between rats cued with the box and rats cued with the box and the light showed that the light cue significantly increased glucose uptake in a neuronal network composed of the lateral, basal, and central nuclei of the amygdala, the anterior and suprachiasmatic hypothalamic nuclei, the nucleus accumbens, the medial septum, and the insular cortex. In contrast, at the 21-day retention interval, both groups demonstrated similar cerebral metabolic activity. The present results indicate that exposure to a light cue increased metabolic activity in the previously mentioned brain structures only when the light acted as an effective retrieval cue, suggesting an involvement of this network in the processes triggered by a retrieval cue. Arguments are provided supporting the notion that the amygdala may play a key role in these processes. Whether the amygdala is a part of a neural network involved in retrieval processes or in neuromodulating systems that favour the efficacy of retrieval processes is also discussed.     

不同情绪背景下来源记忆的ERPs研究*

使用事件相关电位(ERP)技术和测试来源记忆的多键范式,探讨不同情绪效价背景下来源提取的认知神经机制。学习阶段,同时呈现汉字和3种效价情绪图片(重叠);测验阶段,只呈现汉字,要求被试进行四键判断：旧字且背景为中性,旧字且背景为正性,旧字且背景为负性,新字。结果发现：在提取阶段,刺激呈现后300~500 ms,3种情绪背景下来源判断正确项目和来源判断错误项目都比新项目诱发了更正的ERPs(即都存在新旧效应),这一结果反映出刺激呈现后300~500 ms 是一个早期的项目提取阶段,它独立于来源提取。并且,在中性背景下,两种新旧效应没有差别;而在正性和负性背景下,来源判断正确项目的新旧效应显著大于来源判断错误项目。说明相比于中性背景,情绪背景下被试可能更早对来源信息产生熟悉感。在500~650 ms,3种情绪背景下都存在来源判断正确项目的新旧效应,在来源判断错误项目和新项目之间没有显著差异,来源正确与来源错误有显著的新旧效应差异,但二者在头皮分布上是类似的,这反映了晚期的来源提取过程。同时来源正确的新旧效应在这两个时段有显著不同的头皮分布,表明这两个时段有不同加工过程。另外,在500~650 ms,存在显著的情绪效应,正性背景下正确判断来源诱发的ERPs比中性和负性背景下的更正,而且来源判断错误条件下没有情绪效应。综上所述,来源正确和来源错误可能仅仅反映了大脑激活在量上的不同,并不能推断两者存在质的差异;大脑神经活动的早期不仅反映了对记忆项目的熟悉性,而且也受情绪效应的影响,500 ms之后大脑神经活动反映的是对记忆项目回想的过程,这一回想过程也与情绪效应有关,受到情绪效应的调节。     

Prospective memory: the relative effects of encoding, retrieval, and the match between encoding and retrieval

Previous prospective memory studies have revealed some important features of encoding, retrieval, and the match between the encoding and the retrieval that contribute to prospective memory performance. However, these studies have not provided evidence concerning the relative importance of these three factors because no study has investigated all three in a single design. We developed a laboratory-based paradigm that allowed us to manipulate different characteristics of encoding, retrieval, and the match between encoding and retrieval simultaneously in a single experiment. The results of eight experiments showed that all three factors have an influence on prospective memory performance, but that the match between encoding and retrieval has a significantly larger influence than either encoding or retrieval factors.     

Neural correlates of three cognitive processes involved in theory of mind and discourse comprehension

Neuroimaging studies have found that theory of mind (ToM) and discourse comprehension involve similar brain regions. These brain regions may be associated with three cognitive components that are necessarily or frequently involved in ToM and discourse comprehension, including social concept representation and retrieval, domain-general semantic integration, and domain-specific integration of social semantic contents. Using fMRI, we investigated the neural correlates of these three cognitive components by exploring how discourse topic (social/nonsocial) and discourse processing period (ending/beginning) modulate brain activation in a discourse comprehension (and also ToM) task. Different sets of brain areas showed sensitivity to discourse topic, discourse processing period, and the interaction between them, respectively. The most novel finding was that the right temporoparietal junction and middle temporal gyrus showed sensitivity to discourse processing period only during social discourse comprehension, indicating that they selectively contribute to domain-specific semantic integration. Our finding indicates how different domains of semantic information are processed and integrated in the brain and provides new insights into the neural correlates of ToM and discourse comprehension.     

Learning related modulation of functional retrieval networks in man

The medial temporal lobe has been implicated in studies of episodic memory tasks involving spatio-temporal context and object-location conjunctions. We have previously demonstrated that an increased level of practice in a free-recall task parallels a decrease in the functional activity of several brain regions, including the medial temporal lobe, the prefrontal, the anterior cingulate, the anterior insular, and the posterior parietal cortices, that in concert demonstrate a move from elaborate controlled processing towards a higher degree of automaticity. Here we report data from two experiments that extend these initial observations. We used a similar experimental approach but probed for effects of retrieval paradigms and stimulus material. In the first experiment we investigated practice related changes during recognition of object-location conjunctions and in the second during free-recall of pseudo-words. Learning in a neural network is a dynamic consequence of information processing and network plasticity. The present and previous PET results indicate that practice can induce a learning related functional restructuring of information processing. Different adaptive processes likely subserve the functional re-organisation observed. These may in part be related to different demands for attentional and working memory processing. It appears that the role(s) of the prefrontal cortex and the medial temporal lobe in memory retrieval are complex, perhaps reflecting several different interacting processes or cognitive components. We suggest that an integrative interactive perspective on the role of the prefrontal and medial temporal lobe is necessary for an understanding of the processing significance of these regions in learning and memory. It appears necessary to develop elaborated and explicit computational models for prefrontal and medial temporal functions in order to derive detailed empirical predictions, and in combination with an efficient use and development of functional neuroimaging approaches, to further the understanding of the processing significance of these regions in memory.     

Brain activity during the encoding, retention, and retrieval of stimulus representations

Studies of delayed nonmatching-to-sample (DNMS) performance following lesions of the monkey cortex have revealed a critical circuit of brain regions involved in forming memories and retaining and retrieving stimulus representations. Using event-related functional magnetic resonance imaging (fMRI), we measured brain activity in 10 healthy human participants during performance of a trial-unique visual DNMS task using novel barcode stimuli. The event-related design enabled the identification of activity during the different phases of the task (encoding, retention, and retrieval). Several brain regions identified by monkey studies as being important for successful DNMS performance showed selective activity during the different phases, including the mediodorsal thalamic nucleus (encoding), ventrolateral prefrontal cortex (retention), and perirhinal cortex (retrieval). Regions showing sustained activity within trials included the ventromedial and dorsal prefrontal cortices and occipital cortex. The present study shows the utility of investigating performance on tasks derived from animal models to assist in the identification of brain regions involved in human recognition memory.