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Biomedical subjects

Michael D Rugg

Publications and source records attributed to Michael D Rugg.

At least 19 recordsLinked to original sources

Age effects on the neural correlates of episodic retrieval: increased cortical recruitment with matched performance.

Functional neuroimaging investigations have revealed a range of age-related differences in the neural correlates of episodic memory retrieval. Typically, whereas activity is reduced in older compared with younger adults in some regions, other regions are engaged exclusively, or to a greater extent, in older adults. It is unclear whether such differences merely represent the neural correlates of the lower levels of memory performance and impaired recollection typical of older adults. This issue was addressed in the present event-related functional magnetic resonance imaging study. The level of recollection was matched between groups of healthy younger and older adults for a subset of picture items in a source memory task by manipulating the number of study presentations. Contrasts of the activity elicited by old items attracting correct source judgments and correctly identified new items revealed that the 2 groups recruited many of the same brain regions. However, a striking pattern of age-related differences was also observed. In older adults, retrieval-related increases in activity were more widespread and of greater magnitude than in the young. Moreover, regions demonstrating retrieval-related decreases in activity were almost absent in the older participants. These findings suggest an age-related decline in the efficiency with which neural populations support cognitive function.

Adolescent↗

Recollection and the reinstatement of encoding-related cortical activity.

The neural correlates of episodic memory retrieval ("recollection") differ according to the type of information contained in the recollected episode. Such content-specific recollection effects have been hypothesized to reflect the reinstatement of processes or representations active during encoding. Using event-related functional magnetic resonance imaging, we evaluated this hypothesis by directly contrasting the neural activity elicited during the encoding and subsequent recollection of words studied with one of 2 encoding tasks. Study words appearing on pictures of scenes required imagining the word's referent at any location within the scene, whereas words appearing on a blank background required generating a sentence that incorporated the word. On a later memory test, the neural correlates of recollection were operationalized by contrasting the activity elicited during correct "remember" versus "know" responses. Recollected words from the "scene" task elicited activity in regions of left occipital cortex and anterior fusiform gyrus that overlapped regions where encoding-related activity was greater for the scene than sentence task. Conversely, activity elicited by words recollected from the "sentence" task overlapped with a region of ventromedial frontal cortex where encoding-related activity was greater for the sentence task. These content-specific associations between encoding- and recollection-related neural activity strongly support the reinstatement hypothesis of episodic retrieval.

Adolescent↗

Episodic encoding is more than the sum of its parts: an fMRI investigation of multifeatural contextual encoding.

Episodic memories are characterized by their contextual richness, yet little is known about how the various features comprising an episode are brought together in memory. Here we employed fMRI and a multidimensional source memory procedure to investigate processes supporting the mnemonic binding of item and contextual information. Volunteers were scanned while encoding items for which the contextual features (color and location) varied independently, allowing activity elicited at the time of study to be segregated according to whether both, one, or neither feature was successfully retrieved on a later memory test. Activity uniquely associated with successful encoding of both features was identified in the intra-parietal sulcus, a region strongly implicated in the support of attentionally mediated perceptual binding. The findings suggest that the encoding of disparate features of an episode into a common memory representation requires that the features be conjoined in a common perceptual representation when the episode is initially experienced.

Adolescent↗

The relationship between electrophysiological correlates of recollection and amount of information retrieved.

The electrophysiological correlates of recollection were investigated with a modified Remember/Know task in which subjects signaled whether they fully or partially recollected visual object information in each study episode. A positive-going ERP deflection--the left parietal old/new effect--was sensitive to the amount of information recollected, demonstrating greater amplitude when elicited by test items associated with full relative to partial recollection. These findings support prior proposals that the left parietal ERP old/new effect is sensitive to the amount of information recollected from episodic memory. An early-onsetting (ca. 150 ms), left frontal old/new effect differentiated items accorded correct old versus correct new responses regardless of whether the items were endorsed as familiar or recollected. This finding extends the range of circumstances under which early, frontally distributed old/new effects occur, and adds weight to previous suggestions that these effects are a neural correlate of familiarity-driven recognition memory.

Adolescent↗

Neural correlates of differential retrieval orientation: Sustained and item-related components.

Retrieval orientation refers to a cognitive state that biases processing of retrieval cues in service of a specific goal. The present study used a mixed fMRI design to investigate whether adoption of different retrieval orientations - as indexed by differences in the activity elicited by retrieval cues corresponding to unstudied items - is associated with differences in the state-related activity sustained across a block of test trials sharing a common retrieval goal. Subjects studied mixed lists comprising visually presented words and pictures. They then undertook a series of short test blocks in which all test items were visually presented words. The blocks varied according to whether the test items were used to cue retrieval of studied words or studied pictures. In several regions, neural activity elicited by correctly classified new items differed according to whether words or pictures were the targeted material. The loci of these effects suggest that one factor driving differential cue processing is modulation of the degree of overlap between cue and targeted memory representations. In addition to these item-related effects, neural activity sustained throughout the test blocks also differed according to the nature of the targeted material. These findings indicate that the adoption of different retrieval orientations is associated with distinct neural states. The loci of these sustained effects were distinct from those where new item activity varied, suggesting that the effects may play a role in biasing retrieval cue processing in favor of the current retrieval goal.

Adult↗

Electrophysiological dissociation of the neural correlates of recollection and familiarity.

Event-related potentials (ERPs) were employed to investigate electrophysiological correlates of recognition memory in a task that allowed segregation of test items according to whether they were recollected (operationalized by introspective report) or, if recollection failed, their level of familiarity (operationalized by recognition confidence). The amplitude of a negative-going ERP deflection that onsets around 300 ms post-stimulus varied inversely with familiarity strength. This effect was maximal over the left frontal scalp. It did not differ between the ERPs elicited by highly familiar versus recollected items, indicating that the recollection is not merely a consequence of strong familiarity. By contrast, a later positive deflection (onset ca. 500 ms post-stimulus) was enhanced in ERPs elicited by recollected relative to highly familiar items. This effect was maximal over the left posterior scalp and was insensitive to familiarity, as indicated by its absence in the contrast between items judged highly familiar versus highly unfamiliar. The findings constitute a double dissociation between the neural correlates of recollection and familiarity. Together with the results of a parallel functional magnetic resonance imaging study (A.P. Yonelinas et al., J. Neurosci. (2005), 25, 3002-3008), they indicate that recollection and familiarity rely on qualitatively distinct neural systems and strongly support dual-process models of recognition memory.

Adolescent↗

Brain activity before an event predicts later recollection.

Neural activity elicited by an event can predict whether the event is successfully encoded into memory. Here we assessed whether memory encoding relies not only on neural activity that follows an event, but also on activity that precedes it. In two experiments we found that human brain activity elicited by a cue presented just before a word could predict whether the word would be recollected in a later memory test.

Adult↗

Task and content modulate amygdala-hippocampal connectivity in emotional retrieval.

The ability to remember emotional events is crucial for adapting to biologically and socially significant situations. Little is known, however, about the nature of the neural interactions supporting the integration of mnemonic and emotional information. Using fMRI and dynamic models of effective connectivity, we examined regional neural activity and specific interactions between brain regions during a contextual memory retrieval task. We independently manipulated emotional context and relevance of retrieved emotional information to task demands. We show that retrieval of emotionally valenced contextual information is associated with enhanced connectivity from hippocampus to amygdala, structures crucially involved with encoding of emotional events. When retrieval of emotional information is relevant to current behavior, amygdala-hippocampal connectivity increases bidirectionally, under modulatory influences from orbitofrontal cortex, a region implicated in representation of affective value and behavioral guidance. Our findings demonstrate that both memory content and behavioral context impact upon large scale neuronal dynamics underlying emotional retrieval.

Amygdala↗

fMRI correlates of retrieval orientation.

Processes related to the cueing of memory retrieval were investigated using fMRI during a yes/no recognition memory test. Participants studied either pictures or auditory words and were subsequently tested with visual words that either corresponded to studied items (old items) or were unstudied (new items). It was expected that neural activity would differ according to the type of information participants are trying to retrieve: a manifestation of the so-called "retrieval orientation" effect. We replicated robust old/new effects in parietal, prefrontal and anterior medial temporal lobe cortices, which did not differ according to the study material. However, we did observe differential activity to test items in temporoparietal cortex and fusiform cortex as a function of the study material. More specifically, attempts to retrieve words encoded auditorily produced greater activity in auditory cortex than attempts to retrieve words encoded as pictures, whereas the converse was found in fusiform cortex. The above pattern was found for both new and old test items. These findings implicate these regions in constraining the search for specific types of encoded information and thus are in accordance with the transfer-appropriate processing framework. Further, we propose that our results can be seen as an extension of the cortical reinstatement hypothesis, in that the same material-specific cortical regions are engaged during both encoding and retrieval, and this increases the likelihood of successful recognition, or rejection, of retrieval cues in a memory test.

Acoustic Stimulation↗

Modulation of the electrophysiological correlates of retrieval cue processing by the specificity of task demands.

Retrieval orientation refers to the differential processing of retrieval cues according to the type of information sought from memory (e.g., words vs. pictures). In the present study, event-related potentials (ERPs) were employed to investigate whether the neural correlates of differential retrieval orientations are sensitive to the specificity of the retrieval demands of the test task. In separate study-test phases, subjects encoded lists of intermixed words and pictures, and then undertook one of two retrieval tests, in both of which the retrieval cues were exclusively words. In the recognition test, subjects performed 'old/new' discriminations on the test items, and old items corresponded to only one class of studied material (words or pictures). In the exclusion test, old items corresponded to both classes of study material, and subjects were required to respond 'old' only to test items corresponding to a designated class of material. Thus, demands for retrieval specificity were greater in the exclusion test than during recognition. ERPs elicited by correctly classified new items in the two types of test were contrasted according to whether words or pictures were the sought-for material. Material-dependent ERP effects were evident in both tests, but the effects onset earlier and offset later in the exclusion test. The findings suggest that differential processing of retrieval cues, and hence the adoption of differential retrieval orientations, varies according to the specificity of the retrieval goal.

Adolescent↗

ERP correlates of retrieval orientation: direct versus indirect memory tasks.

The neural correlates of the processes involved in cueing memory retrieval were investigated using ERPs to unstudied (new) items in a yes/no recognition test (direct memory task) and a semantic judgement task (indirect memory task). Subjects encoded either pictures or auditory words and were tested on visual words. We replicated previous findings that ERPs to correct rejections of new items in a yes/no recognition test differ according to the study material, with ERPs to words encoded as pictures being more negative (relative to a average mastoid reference) than words encoded auditorily [Hornberger, M., Morcom, A.M., et al., 2004. Neural correlates of retrieval orientation: effects of study-test similarity. J. Cogn. Neurosci. 16 (7), 1196-1210]. This difference was sustained from approximately 450-1,200 ms. An effect of study material on the ERP to new items was found in the indirect memory task but was both earlier onsetting and shorter lived, ca. 250-600 ms. These findings add weight to the concept of 'retrieval orientation' [Rugg, M.D., Wilding, E.L., 2000. Retrieval processing and episodic memory. Trends Cogn. Sci. 4 (3) 108-115] - differential processing of retrieval cues according to the form of the sought-for information - by showing that a putative ERP correlate of retrieval orientation is restricted to direct memory tasks.

Acoustic Stimulation↗

Electrophysiological correlates of retrieval processing: effects of consistent versus inconsistent retrieval demands.

Studies employing event-related potentials (ERPs) during tests of recognition memory have reported differences in neural activity elicited by new test items according to the specific demands of the retrieval task, such as retrieving studied words versus pictures. The present study investigated whether differential processing of new items is possible when retrieval demands vary unpredictably on a trial-by-trial basis. In separate study-test phases, subjects encoded lists of intermixed words and pictures, and undertook retrieval tests with words as test items. Each test item was preceded by a task cue that signaled whether subjects were to attempt to retrieve a word or a picture from the study list. In the "blocked"condition, the targeted study material remained constant throughout the test, whereas in the "mixed"condition, the targeted material varied unpredictably across trials. New-item ERPs were more positive-going when words rather than pictures were targeted in the "blocked" condition, replicating previous findings, but this effect was absent in the "mixed"condition. The findings are consistent with the hypothesis that differential processing of retrieval cues depends upon the adoption of different task sets ("retrieval orientations" that develop over multiple trials and cannot be adjusted merely in response to an instructional cue. Unlike the new-item ERPs, ERPs elicited by the task cues in the mixed condition differed according to targeted material, but only on trials when there was a switch between target material. The implications of these findings for understanding the different retrieval strategies engaged when retrieval demands are consistent versus inconsistent are discussed.

Adolescent↗

MEG source localization under multiple constraints: an extended Bayesian framework.

To use Electroencephalography (EEG) and Magnetoencephalography (MEG) as functional brain 3D imaging techniques, identifiable distributed source models are required. The reconstruction of EEG/MEG sources rests on inverting these models and is ill-posed because the solution does not depend continuously on the data and there is no unique solution in the absence of prior information or constraints. We have described a general framework that can account for several priors in a common inverse solution. An empirical Bayesian framework based on hierarchical linear models was proposed for the analysis of functional neuroimaging data [Friston, K., Penny, W., Phillips, C., Kiebel, S., Hinton, G., Ashburner, J., 2002. Classical and Bayesian inference in neuroimaging: theory. NeuroImage 16, 465-483] and was evaluated recently in the context of EEG [Phillips, C., Mattout, J., Rugg, M.D., Maquet, P., Friston, K., 2005. An empirical Bayesian solution to the source reconstruction problem in EEG. NeuroImage 24, 997-1011]. The approach consists of estimating the expected source distribution and its conditional variance that is constrained by an empirically determined mixture of prior variance components. Estimation uses Expectation-Maximization (EM) to give the Restricted Maximum Likelihood (ReML) estimate of the variance components (in terms of hyperparameters) and the Maximum A Posteriori (MAP) estimate of the source parameters. In this paper, we extend the framework to compare different combinations of priors, using a second level of inference based on Bayesian model selection. Using Monte-Carlo simulations, ReML is first compared to a classic Weighted Minimum Norm (WMN) solution under a single constraint. Then, the ReML estimates are evaluated using various combinations of priors. Both standard criterion and ROC-based measures were used to assess localization and detection performance. The empirical Bayes approach proved useful as: (1) ReML was significantly better than WMN for single priors; (2) valid location priors improved ReML source localization; (3) invalid location priors did not significantly impair performance. Finally, we show how model selection, using the log-evidence, can be used to select the best combination of priors. This enables a global strategy for multiple prior-based regularization of the MEG/EEG source reconstruction.

Bayes Theorem↗

Impairment of specific episodic memory processes by sub-psychotic doses of ketamine: the effects of levels of processing at encoding and of the subsequent retrieval task.

RATIONALE: The precise nature of the impact of the N-methyl-D-aspartate antagonist, ketamine, upon human episodic memory, has yet to be elucidated fully. OBJECTIVES: This study sought to assess the effects of ketamine on the sub-processes facilitating memory encoding and retrieval. METHODS: We evaluated the effects of the drug on a series of memory performance measures depending upon whether it was administered at the encoding or retrieval stage and on the nature of the encoding task used. Twelve healthy volunteers participated in a double-blind, placebo-controlled, randomized, within-subjects study. Intravenous infusions of placebo, 50 ng/ml ketamine or 100 ng/ml ketamine were administered. We investigated the effects of ketamine on three key aspects of episodic memory: encoding vs retrieval processes, source memory, and depth of processing. Data were analysed using both multinomial modelling and standard measures of item discrimination and response bias. RESULTS: Deleterious effects of ketamine on episodic memory were primarily attributable to its effects on encoding, rather than retrieval processes. Recognition memory was impaired for items encoded at an intermediate level of processing, but preserved for shallowly and deeply encoded items. Increased source guessing bias was also observed when encoding took place under ketamine. CONCLUSIONS: The effects of ketamine upon episodic memory seem, therefore, to predominate at encoding. Furthermore, our results are also consistent with a specific impairment of encoding processes that result in subsequent recollective, as opposed to familiarity-based, retrieval. The observed effects are compatible with memory deficits seen in schizophrenia and thus provide some support for the ketamine model of the disease.

Adolescent↗

Encoding and the durability of episodic memory: a functional magnetic resonance imaging study.

Memories vary in their durability even when encoding conditions apparently remain constant. We investigated whether, under these circumstances, memory durability is nonetheless associated with variation in the neural activity elicited during encoding. Event-related functional magnetic resonance imaging data were acquired while volunteers semantically classified visually presented words. Using the "remember/know" procedure, memory for one-half of the words was tested after 30 min and for the remaining half after 48 h. In several regions, including left hippocampus and left dorsal inferior frontal gyrus (IFG), activity at encoding differed depending on whether items were later recollected regardless of study-test delay. Delay-selective effects were also evident, however. Recollection after 48 h was associated with enhanced activity in bilateral ventral IFG, whereas recollection after 30 min was associated with greater fusiform activity. Thus, there is a relationship between the neural activity elicited by an event as it is encoded and the durability of the resulting memory representation.

Adult↗

Separating the brain regions involved in recollection and familiarity in recognition memory.

The neural substrates of recognition memory retrieval were examined in a functional magnetic resonance imaging study designed to separate activity related to recollection from that related to continuous variations in familiarity. Across a variety of brain regions, the neural signature of recollection was found to be distinct from familiarity, demonstrating that recollection cannot be attributed to familiarity strength. In the prefrontal cortex, an anterior medial region was related to recollection, but lateral regions, including the anterior and dorsolateral prefrontal cortex, were related to familiarity. Along the lateral parietal cortex, two functionally distinct regions were also observed: a lateral parietal/temporal region related to recollection and a more superior parietal region involved in familiarity. Similarly, in medial parietal regions, the posterior cingulate was related to recollection, whereas the precuneus was related to familiarity. The hippocampus was related to recollection, but also exhibited an inverse relationship to familiarity-driven recognition confidence. The results indicate that recollection and familiarity rely on different networks of brain regions and provide insights into the functional roles of different regions involved in episodic recognition memory.

Adult↗

Content-specificity of the neural correlates of recollection.

It is widely assumed that episodic retrieval (recollection) involves reinstatement of cortical activity engaged during the processing of an episode when it was initially experienced. It follows from this assumption that the cortical correlates of recollection should differ with the content of what is recollected, and that retrieval of different content should be associated with activity in functionally distinct cortical regions. The present experiment investigated these predictions. Subjects (N=17) studied a mixed list of words and pictures and were then presented with a test list comprised of words only. Test items were studied words, the names of studied pictures, and unstudied (new) words. Functional magnetic resonance images were acquired while the subjects made Remember/Know/New judgments to these test words. Independent of study material, studied items endorsed as Remembered elicited greater activity than correctly classified unstudied items in several regions, including left frontal, left lateral parietal, and posterior cingulate cortex. In addition, Remembered items elicited greater activity in the right hippocampus and parahippocampal gyrus than items accorded Know judgments, replicating previous findings. Analysis of content-specific effects demonstrated a regional double-dissociation within left fusiform cortex; recollected words elicited greater activity than recollected pictures in lateral fusiform, whereas the reverse effect was evident in an anterior fusiform region. The lateral and anterior fusiform areas correspond closely to areas held to be functionally specialized for the processing of visual words and pictures, respectively. Thus, the current findings support the cortical reinstatement hypothesis of episodic retrieval.

Adolescent↗

An empirical Bayesian solution to the source reconstruction problem in EEG.

Distributed linear solutions of the EEG source localisation problem are used routinely. In contrast to discrete dipole equivalent models, distributed linear solutions do not assume a fixed number of active sources and rest on a discretised fully 3D representation of the electrical activity of the brain. The ensuing inverse problem is underdetermined and constraints or priors are required to ensure the uniqueness of the solution. In a Bayesian framework, the conditional expectation of the source distribution, given the data, is attained by carefully balancing the minimisation of the residuals induced by noise and the improbability of the estimates as determined by their priors. This balance is specified by hyperparameters that control the relative importance of fitting and conforming to various constraints. Here we formulate the conventional "Weighted Minimum Norm" (WMN) solution in terms of hierarchical linear models. An "Expectation-Maximisation" (EM) algorithm is used to obtain a "Restricted Maximum Likelihood" (ReML) estimate of the hyperparameters, before estimating the "Maximum a Posteriori" solution itself. This procedure can be considered a generalisation of previous work that encompasses multiple constraints. Our approach was compared with the "classic" WMN and Maximum Smoothness solutions, using a simplified 2D source model with synthetic noisy data. The ReML solution was assessed with four types of source location priors: no priors, accurate priors, inaccurate priors, and both accurate and inaccurate priors. The ReML approach proved useful as: (1) The regularisation (or influence of the a priori source covariance) increased as the noise level increased. (2) The localisation error (LE) was negligible when accurate location priors were used. (3) When accurate and inaccurate location priors were used simultaneously, the solution was not influenced by the inaccurate priors. The ReML solution was then applied to real somatosensory-evoked responses to illustrate the application in an empirical setting.

Algorithms↗