Biomedical subjects

E Tulving

Publications and source records attributed to E Tulving.

At least 19 recordsLinked to original sources

PubMed2004

[Episodic memory: from mind to brain].

Episodic memory is a neurocognitive (brain/mind) system, uniquely different from other memory systems, that enables human beings to remember past experiences. The notion of episodic memory was first proposed some 30 Years ago. At that time it was defined in terms of materials and tasks. It was subsequently refined and elaborated in terms of ideas such as self, subjective time, and autonoetic consciousness. This chapter provides a brief history of the concept of episodic memory, describes how it has changed (indeed greatly changed) since its inception, considers criticisms of it, and then discusses supporting evidence provided by (a) neuropsychological studies of patterns of memory impairment caused by brain damage, and (b) functional neuroimaging studies of patterns of brain activity of normal subjects engaged in various memory tasks. I also suggest that episodic memory is a true, even if as yet generally unappreciated, marvel of nature.

Brain↗

PubMed2001

Episodic memory and common sense: how far apart?

Research has revealed facts about human memory in general and episodic memory in particular that deviate from both common sense and previously accepted ideas. This paper discusses some of these deviations in light of the proceedings of The Royal Society's Discussion Meeting on episodic memory. Retrieval processes play a more critical role in memory than commonly assumed; people can remember events that never happened; and conscious thoughts about one's personal past can take two distinct forms-'autonoetic' remembering and 'noetic' knowing. The serial-dependent-independent (SPI) model of the relations among episodic, semantic and perceptual memory systems accounts for a number of puzzling phenomena, such as some amnesic patients' preserved recognition memory and their ability to learn new semantic facts, and holds that episodic remembering of perceptual information can occur only by virtue of its mediation through semantic memory. Although common sense endows many animals with the ability to remember their past experiences, as yet there is no evidence that humanlike episodic memory-defined in terms of subjective time, self, and autonoetic awareness-is present in any other species.

Animals↗

PubMed2001

Comparative electrophysiological and hemodynamic measures of neural activation during memory-retrieval.

The spatial and temporal characteristics of the brain processes underlying memory retrieval were studied with both event-related potentials (ERP) and positron emission tomography (PET) techniques. Subjects studied lists of 20 words and then performed episodic (old/new judgment) or semantic (living/nonliving decision) retrieval tasks on multiple four-item test lists, each lasting 10 sec. The PET and ERP measurements at test were assessed in relation to both the task (episodic vs. semantic) and the item (old vs. new or living vs. nonliving). Episodic retrieval was associated with increased blood flow in the right frontal lobe (Brodmann Area 10) and a sustained, slowly developing positive ERP shift recorded from the right frontopolar scalp. Semantic retrieval was associated with increased blood flow in the left frontal (Area 45) and temporal (Area 21) lobes but no clear ERP concomitant. The two retrieval tasks also differed from each other in the ERPs to single items in an early (300-500 ms) time window. Item-related comparisons yielded convergent results mainly if the retrieved information was relevant to the given task (e.g., old/new items during episodic retrieval and living/nonliving items during semantic retrieval). Episodically retrieved old items were associated with increased blood flow in the left medial temporal lobe and a transient increase in the amplitude of the late positive component (500-700 ms) of the ERP. Semantically retrieved living items were associated with increased blood flow in the left frontal cortex and anterior cingulate and a transient late frontal slow wave (700-1,500 ms) in the ERPs. These results indicate that the brain regions engaged in memory retrieval are active in either a sustained or transient manner. They map task-related processes to sustained and item-related processes to transient neural activity. But they also suggest that task-related factors can transiently affect early stages of item processing.

Adult↗

PubMed2001

Transperceptual encoding and retrieval processes in memory: a PET study of visual and haptic objects.

An important objective of functional neuroimaging research is to identify neuroanatomical correlates of memory processes such as encoding and retrieval. In typical studies directed at this goal, however, the to-be-remembered information has been presented in a single perceptual modality. Under these conditions it is not known whether the observed brain activity reflects the studied memory process as such or only the memory process in the given modality. The positron emission tomography (PET) study reported here was designed to identify brain regions involved in encoding and retrieval processes specific to visual and haptic modalities, as well as those common to the two modalities. These latter, common regions, were assumed to be associated with "transperceptual" encoding and retrieval processes. Abstract three-dimensional objects, difficult to describe verbally, served as to-be-remembered materials. A multivariate partial least squares analysis of the PET data revealed that transperceptual encoding processes activated right medial temporal lobe, superior prefrontal cortex bilaterally, and posterior inferior temporal gyrus bilaterally. Transperceptual recognition activations were observed in two right orbitofrontal regions and in anterior cingulate. These results provide initial evidence that some processes involved in memory encoding and retrieval operate beyond perceptual processes and in that sense are transperceptual.

Adult↗

PubMed2000

Reactivation of encoding-related brain activity during memory retrieval.

Neuronal models predict that retrieval of specific event information reactivates brain regions that were active during encoding of this information. Consistent with this prediction, this positron-emission tomography study showed that remembering that visual words had been paired with sounds at encoding activated some of the auditory brain regions that were engaged during encoding. After word-sound encoding, activation of auditory brain regions was also observed during visual word recognition when there was no demand to retrieve auditory information. Collectively, these observations suggest that information about the auditory components of multisensory event information is stored in auditory responsive cortex and reactivated at retrieval, in keeping with classical ideas about "redintegration, " that is, the power of part of an encoded stimulus complex to evoke the whole experience.

Adult↗

PubMed2000

Prefrontal cortex and episodic memory retrieval mode.

A multistudy analysis of positron emission tomography data identified three right prefrontal and two left prefrontal cortical sites, as well as a region in the anterior cingulate gyrus, where neuronal activity is correlated with the maintenance of episodic memory retrieval mode (REMO), a basic and necessary condition of remembering past experiences. The right prefrontal sites were near the frontal pole [Brodmann's area (BA) 10], frontal operculum (BA 47/45), and lateral dorsal area (BA 8/9). The two left prefrontal sites were homotopical with the right frontal pole and opercular sites. The same kinds of REMO sites were not observed in any other cerebral region. Many previous functional neuroimaging studies of episodic memory retrieval have reported activations near the frontal REMO sites identified here, although their function has not been clear. Many of these, too, probably have signaled their involvement in REMO. We propose that REMO activations largely if not entirely account for the frontal hemispheric asymmetry of retrieval as described by the original hemispheric encoding retrieval asymmetry model.

Humans↗

PubMed2000

Individual differences in the functional neuroanatomy of verbal discrimination learning revealed by positron emission tomography.

Why do some people have better memory abilities than others? This issue has been of long-standing interest to scientists and lay people. However, using purely behavioral methods, psychologists have made little progress in illuminating it. Now that functional brain imaging techniques have become available to study mind/brain relations, there is a new promise of understanding individual differences in learning and memory in terms of corresponding differences in brain activity. In this paper, we will present a positron emission tomography (PET) study designed to examine individual differences in learning and memory abilities. The basic assumption is that different patterns of brain activity serve as strong predictors of memory performance. Two specific questions were addressed in this study: (i) Can PET illuminate the relations between memory processes and their neuroanatomical correlates among individual learners and rememberers? and (ii) if so, how are these relations affected by the stage of practice on a given memory task? Our PET study examined individual differences in the neuroanatomical correlates of multi-trial verbal discrimination learning in 16 young healthy subjects. The results identified patterns of brain regions in which blood flow correlated with subjects' retrieval performance. However, these regions did not correlate with performance during all learning trials. Instead, a gradual shift was observed from one pattern of brain regions to another over the course of learning. These results suggest that individual differences in memory performance are related to differences in neural activity within specific brain circuits.

Adult↗

PubMed2000

Large scale neurocognitive networks underlying episodic memory.

Large-scale networks of brain regions are believed to mediate cognitive processes, including episodic memory. Analyses of regional differences in brain activity, measured by functional neuroimaging, have begun to identify putative components of these networks. To more fully characterize neurocognitive networks, however, it is necessary to use analytical methods that quantify neural network interactions. Here, we used positron emission tomography (PET) to measure brain activity during initial encoding and subsequent recognition of sentences and pictures. For each type of material, three recognition conditions were included which varied with respect to target density (0%, 50%, 100%). Analysis of large-scale activity patterns identified a collection of foci whose activity distinguished the processing of sentences vs. pictures. A second pattern, which showed strong prefrontal cortex involvement, distinguished the type of cognitive process (encoding or retrieval). For both pictures and sentences, the manipulation of target density was associated with minor activation changes. Instead, it was found to relate to systematic changes of functional connections between material-specific regions and several other brain regions, including medial temporal, right prefrontal and parietal regions. These findings provide evidence for large-scale neural interactions between material-specific and process-specific neural substrates of episodic encoding and retrieval.

Adult↗

PubMed1999

Task-related and item-related brain processes of memory retrieval.

In all cognitive tasks, general task-related processes operate throughout a given task on all items, whereas specific item-related processes operate differentially on individual items. In typical functional neuroimaging experiments, these two sets of processes have usually been confounded. Herein we report a combined positron emission tomography and event-related potential (ERP) experiment that was designed to distinguish between neural correlates of task-related and item-related processes of memory retrieval. Two retrieval tasks, episodic and semantic, were crossed with episodic (old/new) and semantic (living/nonliving) properties of individual items to yield evidence of regional brain activity associated with task-related processes, item-related processes, and their interaction. The results showed that episodic retrieval task was associated with increased blood flow in right prefrontal and posterior cingulate cortex, as well as with a sustained right-frontopolar-positive ERP, but that the semantic retrieval task was associated with left frontal and temporal lobe activity. Retrieval of old items was associated with increased blood flow in the left medial temporal lobe and with a brief late positive ERP component. The results provide converging hemodynamic and electrophysiological evidence for the distinction of task- and item-related processes, show that they map onto spatially and temporally distinct patterns of brain activity, and clarify the hemispheric encoding/retrieval asymmetry (HERA) model of prefrontal encoding and retrieval asymmetry.

Adult↗

PubMed1999

Positron emission tomography correlations in and beyond medial temporal lobes.

This article discusses the potential usefulness of brain/ behavior correlational analyses in functional neuroimaging studies of memory, and how such analyses can illuminate the role of medial temporal lobes (MTL) and the hippocampus in episodic and declarative memory processes such as encoding and retrieval. Reanalysis of the results of four previously reported positron emission tomography (PET) studies yielded evidence of both positive and negative between-subjects correlations between recognition-memory accuracy and regional blood flow. The sites of these correlations were in MTL regions as well as in other cortical and subcortical areas, including frontal lobes (Brodmann areas 6, 9, 10, 11, and 47), temporal lobes (BAs 21, 22, and 38), insula, fusiform gyrus, and cuneus/precuneus. These findings were discussed with respect to issues such as localization of the correlation sites, the distinction between brain sites revealed by brain/cognition correlational analyses ("how" sites) and those yielded by cognitive subtraction methods ("what" sites), the tendency of the "how" sites in MTL to occur in the left hemisphere, the tendency of other "how" sites to occur in one or the other hemisphere, rather than bilaterally, and the meaning and "reality" of both brain/behavior correlations and task-related activations. Because of the known incidence of false-positives, all neuroimaging data, including those involving the localization of "what" and "how" memory sites in MTL and other brain regions, need to be interpreted cautiously, and findings of individual studies should not be overinterpreted.

Brain↗

PubMed1998

Episodic and declarative memory: role of the hippocampus.

The fact that medial temporal lobe structures, including the hippocampus, are critical for declarative memory is firmly established by now. The understanding of the role that these structures play in declarative memory, however, despite great efforts spent in the quest, has eluded investigators so far. Given the existing scenario, novel ideas that hold the promise of clarifying matters should be eagerly sought. One such idea was recently proposed by Vargha-Khadem and her colleagues (Science 1997; 277:376-380) on the basis of their study of three young people suffering from anterograde amnesia caused by early-onset hippocampal pathology. The idea is that the hippocampus is necessary for remembering ongoing life's experiences (episodic memory), but not necessary for the acquisition of factual knowledge (semantic memory). We discuss the reasons why this novel proposal makes good sense and why it and its ramifications should be vigorously pursued. We review and compare declarative and episodic theories of amnesia, and argue that the findings reported by Vargha-Khadem and her colleagues fit well into an episodic theory that retains components already publicized, and adds new ones suggested by the Vargha-Khadem et al. study. Existing components of this theory include the idea that acquisition of factual knowledge can occur independently of episodic memory, and the idea that in anterograde amnesia it is quite possible for episodic memory to be more severely impaired than semantic memory. We suggest a realignment of organization of memory such that declarative memory is defined in terms of features and properties that are common to both episodic and semantic memory. The organization of memory thus modified gives greater precision to the Vargha-Khadem et al. neuroanatomical model in which declarative memory depends on perihippocampal cortical regions but not on the hippocampus, whereas episodic memory, which is separate from declarative memory, depends on the hippocampus.

Amnesia↗

PubMed1998

Hippocampal PET activations of memory encoding and retrieval: the HIPER model.

A meta-analysis of experimentally induced changes in blood flow ("activations") in positron emission tomography (PET) studies of memory has revealed an orderly functional anatomic pattern: Activations in the hippocampal region associated with episodic memory encoding are located primarily in the rostral portions of the region, whereas activations associated with episodic memory retrieval are located primarily in the caudal portions. These findings are based on an analysis of a sample of 54 "hippocampal encoding and retrieval" activations that were culled from an overall database consisting of 52 published PET studies of memory. We refer to this general pattern of rostrocaudal gradient of encoding and retrieval PET activations as the HIPER (Hippocampal Encoding/Retrieval) model. The model suggests a division of memory-related labor between the rostral and caudal portions of the hippocampal formation. Because functional anatomic pattern of encoding and retrieval activation that defines the HIPER model was unprecedented and unexpected, it is difficult to relate the model to what is already known or thought about functional neuroanatomy of episodic memory in the hippocampal regions. The model is interesting primarily because its exploration may yield fresh insights into the neural basis of human memory.

Brain Mapping↗

PubMed1998

Functional brain imaging of episodic and semantic memory with positron emission tomography.

Human memory is composed of several independent but interacting systems. These include a system for remembering general knowledge, semantic memory, and a system for recollection of personal events, episodic memory. The results of positron emission tomography (PET) studies of regional cerebral blood flow indicate that networks of distributed brain regions subserve episodic and semantic memory. Some networks seem to be generally engaged in memory processes whereas the involvement of others is specific to factors such as the type of information to be remembered or the level of retrieval success. The PET findings help to understand memory dysfunction (a) by showing that multiple brain regions are involved in different memory processes and (b) by sharpening the interpretation of the functional role of different brain regions.

Brain↗

PubMed1998

Unilateral medial temporal lobe memory impairment: type deficit, function deficit, or both?

Previous research has characterized memory deficits resulting from unilateral hippocampal system damage as 'material specific', suggesting that left damage results in verbal memory impairment with preservation of visuospatial function and the converse with right damage. Implicit within this hypothesis are the assumptions that the systems are independent and memory is lateralized for each type of material. To test the verbal component of this hypothesis, unilateral hippocampal lesion and commissurotomy patients were compared with controls on a multiple-list free-recall task. The material specific hypothesis predicts severe impairment only with left lesions; right lesions and commissurotomy patients should be only minimally impaired. However, secondary memory was compromised at immediate recall for all patient groups, with both unilateral groups showing comparable and severe verbal episodic memory deficits. Final testing across all lists also revealed severe impairment in commissurotomy patients. Finding both unilateral groups to be similarly impaired for verbal material is taken as evidence against a material specific deficit during this verbal episodic memory task. Although previous data suggest that left patients are considerably more impaired during some verbal tasks, this may not be specific to the material, but rather the combination of material and task demands. Implications for the material specific hypothesis are discussed.

Adult↗

PubMed1998

Episodic memory and the self in a case of isolated retrograde amnesia.

Isolated retrograde amnesia is defined as impaired recollection of experiences pre-dating brain injury with relatively preserved anterograde learning and memory. We present findings from a patient (M.L.) with isolated retrograde amnesia following severe traumatic brain injury (TBI) that address hypotheses of the interrelationships of focal neuropathology, episodic memory and the self. M.L. is densely amnesic for experiences predating his injury, but shows normal anterograde memory performance on a variety of standard tests of recall and recognition. The cognitive processes underlying this performance were examined with the remember/know technique, which permits separation of episodic from non-episodic contributions to memory tests by quantifying subjects' reports of re-experiencing aspects of the encoding episode. The results demonstrated that M.L. does not episodically re-experience post-injury events to the same extent as control subjects, although he can use familiarity or other non-episodic processes to distinguish events he has experienced from those he has not experienced. M.L.'s MRI showed damage to the right ventral frontal cortex and underlying white matter, including the uncinate fasciculus, a frontotemporal band of fibres previously hypothesized to mediate retrieval of specific events from one's personal past. Recent functional neuroimaging evidence of an association between right frontal lobe functioning and episodic retrieval demands suggest that M.L.'s memory deficits are related to this focal injury. This hypothesis was supported by right frontal polar hypoactivation in M.L. in response to episodic retrieval demands when he was examined with a cognitive activation H2(15)O PET paradigm that reliably activated this frontal region in both healthy controls and patients with TBI carefully matched to M.L. (but without isolated retrograde amnesia). He also showed increased left inferomedial temporal activation relative to control subjects, suggesting that his spared anterograde memory is mediated through increased reliance on medial temporal lobe structures. Re-experiencing events as part of one's past is based on autonoetic awareness, i.e. awareness of oneself as a continuous entity across time. This form of awareness also supports the formulation of future goals and the implementation of a behavioural guidance system to achieve them. The findings from this study converge to suggest that M.L. has impaired autonoetic awareness attributable to right ventral frontal lobe injury, including right frontal-temporal disconnection. Reorganized brain systems mediate certain preserved cognitive operations in M.L., but without the normal complement of information concerning the self with respect to both past and future events.

Adult↗

PubMed1997

Age-related differences in effective neural connectivity during encoding and recall.

Age-related differences in brain activity may reflect local neural changes in the regions involved or they may reflect a more global transformation of brain function. To investigate this issue, we applied structural equation modeling to the results of a positron emission tomography (PET) study in which young and old adults encoded and recalled word pairs. In the young group there was a shift from positive interactions involving the left prefrontal cortex during encoding to positive interactions involving the right prefrontal cortex during recall, whereas in the old group frontal interactions were mixed during encoding and bilaterally positive during recall. The present results suggest that age-related changes in neural activation are partly due to age-related changes in effective connectivity in the neural network underlying the task.

Adult↗

PubMed1997

Event-related brain potential correlates of two states of conscious awareness in memory.

We report an event-related potential (ERP) experiment of human recognition memory that explored the relation between conscious awareness and electrophysiological activity of the brain. We recorded ERPs from healthy adults while they made "remember" and "know" recognition judgments about previously seen words. These two kinds of judgments reflect "autonoetic" and "noetic" awareness, respectively. The ERP effects differed between the two kinds of awareness while they were similar for "true" and "false" recognition. Noetic awareness was associated with a temporoparietal positivity in the N400 range (325-600 ms) and a late (600-1,000 ms) frontocentral negativity, whereas autonoetic awareness was associated with a widespread, late, bifrontal and left parietotemporal (600-1000 ms) positivity. In the very late (1,300-1, 900 ms) time window, a right frontal positivity was observed for both remember and know judgments of both true and false targets. These results provide physiological evidence for two types of conscious awareness in episodic memory retrieval.

Adult↗

PubMed1997

Age-related differences in neural activity during memory encoding and retrieval: a positron emission tomography study.

Positron emission tomography (PET) was used to compare regional cerebral blood flow (rCBF) in young (mean 26 years) and old (mean 70 years) subjects while they were encoding, recognizing, and recalling word pairs. A multivariate partial-least-squares (PLS) analysis of the data was used to identify age-related neural changes associated with (1) encoding versus retrieval and (2) recognition versus recall. Young subjects showed higher activation than old subjects (1) in left prefrontal and occipito-temporal regions during encoding and (2) in right prefrontal and parietal regions during retrieval. Old subjects showed relatively higher activation than young subjects in several regions, including insular regions during encoding, cuneus/precuneus regions during recognition, and left prefrontal regions during recall. Frontal activity in young subjects was left-lateralized during encoding and right-lateralized during recall [hemispheric encoding/retrieval asymmetry (HERA)], whereas old adults showed little frontal activity during encoding and a more bilateral pattern of frontal activation during retrieval. In young subjects, activation in recall was higher than that in recognition in cerebellar and cingulate regions, whereas recognition showed higher activity in right temporal and parietal regions. In old subjects, the differences in blood flow between recall and recognition were smaller in these regions, yet more pronounced in other regions. Taken together, the results indicate that advanced age is associated with neural changes in the brain systems underlying encoding, recognition, and recall. These changes take two forms: (1) age-related decreases in local regional activity, which may signal less efficient processing by the old, and (2) age-related increases in activity, which may signal functional compensation.

Adult↗