Biomedical subjects

E Tulving

Publications and source records attributed to E Tulving.

At least 37 records · Page 2Linked to original sources

PubMed1997

Cognitive subtractions may not add up: the interaction between semantic processing and response mode.

Determining the areas of brain activity associated with cognitive processing has typically relied on the use of a subtraction paradigm, which is based on the premise that the neural processes underlying behavior are additive. If the additivity assumption is valid then brain regions associated with a semantic processing task should be the same regardless of how participants make a response. To investigate this proposition, participants underwent six PET scans, in which they made semantic or letter word judgments, responding "yes" or "no" in three different modes: mouse-clicking, spoken response, or silent thought. Analyses showed an increase in regional cerebral blood flow associated with semantic processing in the left inferior frontal cortex, anterior cingulate, and right cerebellum for all three response conditions. However, there was a significant interaction: the greatest increase was observed in the mouse-click condition and the weakest change seen with silent thought. Moreover, other areas of the brain were uniquely activated for each response mode. The results indicate that different areas of the brain were recruited for semantic processing depending on how participants had to organize their responses. Implications for the additivity assumption and methods of analysis to be used in conjunction with the subtraction technique are discussed.

Adult↗

PubMed1997

Brain regions differentially involved in remembering what and when: a PET study.

Recollecting a past episode involves remembering not only what happened but also when it happened. We used positron emission tomography (PET) to directly contrast the neural correlates of item and temporalorder memory. Subjects studied a list of words and were then scanned while retrieving information about what words were in the list or when they occurred within the list. Item retrieval was related to increased neural activity in medial temporal and basal forebrain regions, whereas temporal-order retrieval was associated with activations in dorsal prefrontal, cuneus/precuneus, and right posterior parietal regions. The dissociation between temporal and frontal lobe regions confirms and extends previous lesion data. The results show that temporal-order retrieval involves a network of frontal and posterior brain regions.

Adult↗

PubMed1997

Memory beyond the hippocampus.

Improved neuroanatomical knowledge, technical and methodological innovations (such as PET), and more refined conceptualizations of memory have inspired a reappraisal of theoretical beliefs regarding the role of the hippocampus in memory. In the past few years, it has become apparent that the influence of the medial temporal lobe regions extends beyond memory and that memory processes (such as encoding, consolidation and retrieval) involve not only the hippocampus and the medial temporal and diencephalic regions, but also widely distributed neocortical and perhaps even cerebellar regions.

Animals↗

PubMed1997

Toward a theory of episodic memory: the frontal lobes and autonoetic consciousness.

Adult humans are capable of remembering prior events by mentally traveling back in time to re-experience those events. In this review, the authors discuss this and other related capabilities, considering evidence from such diverse sources as brain imaging, neuropsychological experiments, clinical observations, and developmental psychology. The evidence supports a preliminary theory of episodic remembering, which holds that the prefrontal cortex plays a critical, supervisory role in empowering healthy adults with autonoetic consciousness-the capacity to mentally represent and become aware of subjective experiences in the past, present, and future. When a rememberer mentally travels back in subjective time to re-experience his or her personal past, the result is an act of retrieval from episodic memory.

Adult↗

PubMed1996

General and specific brain regions involved in encoding and retrieval of events: what, where, and when.

Remembering an event involves not only what happened, but also where and when it occurred. We measured regional cerebral blood flow by positron emission tomography during initial encoding and subsequent retrieval of item, location, and time information. Multivariate image analysis showed that left frontal brain regions were always activated during encoding, and right superior frontal regions were always activated at retrieval. Pairwise image subtraction analyses revealed information-specific activations at (i) encoding, item information in left hippocampal, location information in right parietal, and time information in left fusiform regions; and (ii) retrieval, item in right inferior frontal and temporal, location in left frontal, and time in anterior cingulate cortices. These results point to the existence of general encoding and retrieval networks of episodic memory whose operations are augmented by unique brain areas recruited for processing specific aspects of remembered events.

Adult↗

PubMed1996

Network analysis of positron emission tomography regional cerebral blood flow data: ensemble inhibition during episodic memory retrieval.

Two important objectives in the neuroscience of memory are (1) identification of neural pathways involved in memory processes; and (2) characterization of the pattern of interactions between these pathways. Functional neuroimaging can contribute to both of these goals. Using image subtraction analysis of regional cerebral blood flow data measured with positron emission tomography, we identified brain regions that changed activity during episodic memory retrieval (visual work recognition). Relative to a baseline reading task, decreased activity was observed in bilateral prefrontal, bilateral anterior and posterior temporal, and posterior cingulate cortices. Brain regions showing increased activity were the right prefrontal (different from deactivated regions), left anterior cingulate, and left occipital cortices, and vermis of cerebellum. We then performed a network analysis with structural equation modeling to test the hypothesis that regional decreases came about through active inhibition by regions showing increased activity during retrieval. This analysis demonstrated that the influence of activated regions on deactivated regions was more negative during retrieval than during reading, confirming the inhibition hypothesis. Such confirmation could not have been made from the subtraction analysis alone because decreases can come about, at the very least, through reduction of functional influences as well as by active inhibition. The concepts of ensemble excitation and inhibition, as defined through network analysis, are introduced. We argue that is is critical to examine the combined pattern of excitatory and inhibitory influences to fully appreciate the neural basis of episodic memory.

Analysis of Variance↗

PubMed1996

Activation of medial temporal structures during episodic memory retrieval.

Medial temporal lobe structures have been implicated in human episodic memory. Patients with medial temporal lesions show memory deficits, and functional neuroimaging studies have revealed activation in this region during episodic encoding and retrieval when data are averaged over a sample of subjects. The relevance of such observations for memory performance has remained unclear, however. Here we have used positron emission tomography (PET) to examine cerebral blood flow related to verbal episodic retrieval. We observed strong positive correlations between retrieval and blood flow in left medial temporal structures in individual normal human subjects. In addition, multivariate analysis showed that regions in the left medial temporal lobe were dominant components of a pattern of brain regions that distinguished a high-retrieval condition from conditions of lower retrieval. These results suggest that medial temporal activity is related to retrieval success rather than retrieval attempt, possibly by reflecting reactivation of stored patterns.

Brain Mapping↗

PubMed1996

The neural correlates of intentional learning of verbal materials: a PET study in humans.

The purpose of this study was to identify the brain regions invoked when subjects attempt to learn verbal materials for a subsequent memory test. Twelve healthy subjects undertook two different tasks: reading and encoding of word pairs, while they were being scanned using [15O]H2O positron emission tomography (PET). As expected, the encoding pairs were remembered much better (recall 39% vs. 8%; P < 0.001) than reading pairs in a subsequent memory test. The encoding scans, as compared to reading scans, showed activation of the left prefrontal cortex, the anterior cingulate cortex and the left medial temporal cortex. The left prefrontal activations were in two discrete regions: (i) a left anterior and inferior left prefrontal (Brodmann's areas 45, 46) which we attribute to semantic processing; and (ii) a left posterior mid-frontal region (BA 6, 44) which may reflect rote rehearsal. We interpret the data to suggest that when subjects use cognitive strategies of semantic processing and rote-rehearsal to learn words, they invoke discrete regions of the left prefrontal cortex. And this activation of the left prefrontal cortex along with the medial temporal region leads to a neurophysiological memory trace which can be used to guide subsequent memory retrieval.

Adult↗

PubMed1995

Functional brain maps of retrieval mode and recovery of episodic information.

Positron emission tomography (PET) was used to identify brain regions associated with two component processes of episodic retrieval; those related to thinking back in subjective time (retrieval mode) and those related to actual recovery of stored information (ecphory). Healthy young subjects recognized words that had been encoded with respect to meaning or the speaker's voice. Regardless of how the information had been encoded, recognition was associated with increased activation in regions in right prefrontal cortex, left anterior cingulate, and cerebellum. These activations reflect retrieval mode. Recognition following meaning encoding was specifically associated with increased activation in left temporal cortex, and recognition following voice encoding involved regions in right orbital frontal and parahippocampal cortex. These activations reflect ecphory of differentially encoded information.

Adult↗

PubMed1995

Functional role of the prefrontal cortex in retrieval of memories: a PET study.

Retrieval of information from episodic memory involves the processes invoked by the attempt to remember (retrieval attempt) as well as processes associated with the successful retrieval of stored information (ecphory). Previous PET studies of memory have shown an activation of the prefrontal cortex in memory retrieval tasks, and we hypothesised that this activation represents retrieval attempt, not ecphory. This hypothesis was directly directed using [15O]H2 PET imaging in 19 healthy subjects who performed three matched tasks which involved different levels of retrieval attempt and ecphory. The results showed that retrieval attempt was associated with activation of the prefrontal cortex, right greater than left, while ecphory involved the posterior cortical regions. These findings illuminate the functional role of the different neuroanatomical regions involved in episodic remembering.

Adult↗

PubMed1995

Cognitive processes and cerebral cortical fundi.

Human brain evolution has resulted in a large increase in cortical folding as a result of which 60% of the cerebral cortical mantle is buried within sulci. Cortical regions within the sulci, and especially in the fundal zones (fundi) at the bottom of sulci, differ from the rest of the cortex in a number of ways with respect to anatomical and histological morphology. Although physiological implications of the fundal morphology have been discussed from time to time, and although scattered evidence hints at a special functional role for fundi, until recently there have been few empirical facts to guide the inquiry into a possibly special physiological function of fundal zones. In this article we review findings yielded by positron emission tomography studies showing that the peaks of changes in neuronal activity are frequently observed in and near fundi. We discuss, but do not accept, the possibility that these findings reflect either the partial volume effect or the course of cerebral blood vessels. Instead, because of a coarse correlation observed between fundal fraction (the proportion of fundally related activity peaks) and the apparent cognitive complexity of the tasks probed, and in light of the anatomical evidence reviewed, we propose the hypothesis that cortical sulcal and fundal regions play a distinctive role in higher cognitive processing.

Brain Mapping↗

PubMed1995

Frontal lobe damage produces episodic memory impairment.

This article reports the outcome of a meta-analysis of the relation between the frontal lobes and memory as measured by tests of recognition, cued recall, and free recall. We reviewed experiments in which patients with documented, circumscribed frontal pathology were compared with normal control subjects on these three types of tests. Contrary to conventional wisdom, there is strong evidence that frontal damage disrupts performance on all three types of tests, with the greatest impairment in free recall, and the smallest in recognition.

Amnesia↗

PubMed1994

Novelty encoding networks in the human brain: positron emission tomography data.

Data from positron emission tomography (PET) studies showed novelty activations--higher regional cerebral blood flow associated with perceiving novel rather than familiar stimuli. Regions in the right 'expanded' limbic system--hippocampal formation, parahippocampal gyrus, retrosplenial cortex, thalamus, subcallosal area, the border between cortical areas 32 and 10, anterior and inferior cingulate cortex, putamen, and medial prefrontal cortex--showed such activations for complex pictures. Because novel information is usually encoded for storage in memory, these regions can be seen as constituting components of a visual/spatial novelty encoding network. Insular, opercular and temporal regions (e.g. area 37) showed novelty activations not only for visual pictures but also for auditorily presented sentences, and can be thought of as components of a transmodal novelty encoding network.

Auditory Pathways↗

PubMed1994

The role of the left prefrontal cortex in verbal processing: semantic processing or willed action?

This study was designed to test the various proposed explanations (semantic processing, willed action, production of a spoken response) for the unilateral activation of the left prefrontal cortex noted in PET studies of verbal processing. Twenty subjects underwent 15O-water PET scans while undertaking a lexical task (detecting the letter 'a' in visually presented words) and a semantic task (categorizing nouns into living/non-living). The semantic task resulted in a significant unilateral left dorsolateral prefrontal activation. This finding suggests that the left inferior prefrontal cortex is the anatomical region involved in 'working with meaning', and that the activation does not reflect willed action, is not task-specific and is not attributable to the requirements of a spoken response.

Adult↗

PubMed1994

Cognitive processes and cerebral cortical fundi: findings from positron-emission tomography studies.

Positron-emission tomography (PET) studies of regional cerebral blood flow have provided evidence relevant to localization of cognitive functions. The critical loci identified in these studies are typically described in terms of macroanatomically labeled cortical and subcortical regions. We report the results of a meta-analysis of localization of changes in blood flow, based on nearly 1000 cerebral cortical peaks of activity obtained from groups of subjects in 30 PET studies. The results showed that, on average, 47% of these peaks were localized within the fundus regions of cortical sulci. This is an unexpectedly high proportion because fundal regions compose < 8% of the cortical mantle. Further analysis suggested a coarse correlation between the extent of fundal activation observed in different studies and the estimated cognitive complexity of the tasks used in the studies. These findings are potentially interesting because (i) the preponderance of fundal activation has implications for the interpretation of the PET data, (ii) they suggest that cortical sulcal and fundal regions may play a distinctive role in higher cognitive processing, or (iii) both of the above.

Attention↗

PubMed1994

Neuroanatomical correlates of encoding in episodic memory: levels of processing effect.

Cognitive studies of memory processes demonstrate that memory for stimuli is a function of how they are encoded; stimuli processed semantically are better remembered than those processed in a perceptual or shallow fashion. This study investigates the neural correlates of this cognitive phenomenon. Twelve subjects performed two different cognitive tasks on a series of visually presented nouns. In one task, subjects detected the presence or absence of the letter a; in the other, subjects categorized each noun as living or nonliving. Positron emission tomography (PET) scans using 15O-labeled water were obtained during both tasks. Subjects showed substantially better recognition memory for nouns seen in the living/nonliving task, compared to nouns seen in the a-checking task. Comparison of the PET images between the two cognitive tasks revealed a significant activation in the left inferior prefrontal cortex (Brodmann's areas 45, 46, 47, and 10) in the semantic task as compared to the perceptual task. We propose that memory processes are subserved by a wide neurocognitive network and that encoding processes involve preferential activation of the structures in the left inferior prefrontal cortex.

Adult↗

PubMed1994

Neuroanatomical correlates of retrieval in episodic memory: auditory sentence recognition.

This study used positron emission tomography (PET) to investigate the neuroanatomical correlates of remembering previously experienced events. Twelve young healthy adults listened to "old" meaningful sentences which they had studied 24 hr previously. As a control task the subjects listened to comparable "new" sentences that they had never heard before. Regional cerebral blood flow associated with each task was measured by PET scans using 15O-labeled water. Comparison (old-sentence task minus new-sentence task) of the PET images revealed an extended strip of increased blood flow in the right dorsolateral prefrontal cortex (Brodmann's areas 10, 46, and 9) and the anterior portion of area 6. Other principal regions of increased blood flow were situated around the left anterior cingulate sulcus and bilaterally in the parietal lobes (areas 7 and 40). Major decreases in blood flow were situated bilaterally in the temporal lobes (areas 21, 22, 41, and 42). A high proportion of activity changes seemed to be located in the depths of cortical sulci. Increases in blood flow are seen as reflecting the operations of a widely distributed neuronal network involving prefrontal and parietal cortical regions that subserves the conscious recollection of previously experienced events. Decreases in blood flow in the temporal auditory areas are interpreted as reflecting auditory priming. The prevalence of sulcal blood-flow changes may reflect extensive cortical gyrification; it may also indicate that memory-related processes rely on the densely packed neuropil of sulcal regions.

Adult↗

PubMed1994

Hemispheric encoding/retrieval asymmetry in episodic memory: positron emission tomography findings.

Data are reviewed from positron emission tomography studies of encoding and retrieval processes in episodic memory. These data suggest a hemispheric encoding/retrieval asymmetry model of prefrontal involvement in encoding and retrieval of episodic memory. According to this model, the left and right prefrontal lobes are part of an extensive neuronal network that subserves episodic remembering, but the two prefrontal hemispheres play different roles. Left prefrontal cortical regions are differentially more involved in retrieval of information from semantic memory and in simultaneously encoding novel aspects of the retrieved information into episodic memory. Right prefrontal cortical regions, on the other hand, are differentially more involved in episodic memory retrieval.

Brain↗