Biomedical subjects

Michael D Rugg

Publications and source records attributed to Michael D Rugg.

At least 19 recordsLinked to original sources

PubMed2006

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↗

PubMed2006

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↗

PubMed2006

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↗

PubMed2006

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↗

PubMed2006

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↗

PubMed2006

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↗

PubMed2005

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↗

PubMed2005

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↗

PubMed2005

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↗

PubMed2005

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↗

PubMed2005

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↗

PubMed2005

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↗

PubMed2005

Further dissociating the processes involved in recognition memory: an FMRI study.

Based on an event-related potential study by Rugg et al. [Dissociation of the neural correlates of implicit and explicit memory. Nature, 392, 595-598, 1998], we attempted to isolate the hemodynamic correlates of recollection, familiarity, and implicit memory within a single verbal recognition memory task using event-related fMRI. Words were randomly cued for either deep or shallow processing, and then intermixed with new words for yes/no recognition. The number of studied words was such that, whereas most were recognized ("hits"), an appreciable number of shallow-studied words were not ("misses"). Comparison of deep hits versus shallow hits at test revealed activations in regions including the left inferior parietal gyrus. Comparison of shallow hits versus shallow misses revealed activations in regions including the bilateral intraparietal sulci, the left posterior middle frontal gyrus, and the left frontopolar cortex. Comparison of hits versus correct rejections revealed a relative deactivation in an anterior left medial-temporal region (most likely the perirhinal cortex). Comparison of shallow misses versus correct rejections did not reveal response decreases in any regions expected on the basis of previous imaging studies of priming. Given these and previous data, we associate the left inferior parietal activation with recollection, the left anterior medial-temporal deactivation with familiarity, and the intraparietal and prefrontal responses with target detection. The absence of differences between shallow misses and correct rejections means that the hemodynamic correlates of implicit memory remain unclear.

Adolescent↗

PubMed2005

Electrophysiological dissociation of picture versus word encoding: the distinctiveness heuristic as a retrieval orientation.

Event-related potentials (ERPs) were used to investigate the neural processes underlying the distinctiveness heuristic-a response mode in which participants expect to remember vivid details of an experience and make recognition decisions based on this metacognitive expectation. One group of participants studied pictures and auditory words; another group studied visual and auditory words. Studied and novel items were presented at test as words only, with all novel items repeating after varying lags. ERP differences were seen between the word and picture groups for both studied and novel items. For the novel items, ERP differences were largest in frontal and central midline electrodes. In separate analyses, the picture group showed the greatest ERP differences between item types in a parietally based component from 550 to 1000 msec, whereas the word group showed the greatest differences in a frontally based component from 1000 to 2000 msec. The authors suggest that the distinctiveness heuristic is a retrieval orientation that facilitates reliance upon recollection to differentiate between item types. Although the picture group can use this heuristic and its retrieval orientation on the basis of recollection, the word group must engage additional postretrieval processes to distinguish between item types, reflecting the use of a different retrieval orientation.

Adult↗

PubMed2005

Effects of divided attention on fMRI correlates of memory encoding.

Performing a secondary task concurrently with a study task has a detrimental effect on later memory for studied items. To investigate the mechanisms underlying this effect, the processing resources available for an incidental encoding task were varied by manipulating secondary task difficulty. fMRI data were acquired as volunteers (n = 16) made animacy decisions to visually presented study words while concurrently performing either an easy or a hard auditory monitoring task. "Subsequent memory effects"--greater activity at study for words later remembered versus words later forgotten--were identified in the left ventral inferior frontal gyrus and the left anterior hippocampus. These effects did not vary according to whether the encoding task was performed concurrently with the easy or the hard secondary task. However, as secondary task difficulty increased, study-item activity declined and auditory-item activity increased in dorsolateral prefrontal and superior parietal regions that have been implicated in the support of executive and control functions. The findings suggest that dividing attention during encoding influences the probability of engaging the encoding operations that support later episodic memory, but does not alter the nature of the operations themselves. The findings further suggest that the probability of engaging these encoding operations depends on the level of general processing resources engaged in service of the study task.

Adult↗

PubMed2004

Remembrance of odors past: human olfactory cortex in cross-modal recognition memory.

Episodic memory is often imbued with multisensory richness, such that the recall of an event can be endowed with the sights, sounds, and smells of its prior occurrence. While hippocampus and related medial temporal structures are implicated in episodic memory retrieval, the participation of sensory-specific cortex in representing the qualities of an episode is less well established. We combined functional magnetic resonance imaging (fMRI) with a cross-modal paradigm, where objects were presented with odors during memory encoding. We then examined the effect of odor context on neural responses at retrieval when these same objects were presented alone. Primary olfactory (piriform) cortex, as well as anterior hippocampus, was activated during the successful retrieval of old (compared to new) objects. Our findings indicate that sensory features of the original engram are preserved in unimodal olfactory cortex. We suggest that reactivation of memory traces distributed across modality-specific brain areas underpins the sensory qualities of episodic memories.

Adolescent↗

PubMed2004

An event-related potential study of two kinds of source judgment errors.

The neural correlates of source judgment errors for confusable and nonconfusable source-item pairings were investigated. At study, subjects were presented with a series of lists of word pairs. Word pairs belonging to the same list were formed by pairing one of two associated words with different associates of an unstudied theme word (e.g., ship-frost, captain-chilly, ship-freeze, captain-frigid, etc., in one list; wife-pane, husband-glass, wife-curtain, husband-sill, etc., in another list). At test, subjects discriminated 'Old pairs' (e.g., ship-frost) from 'Intra' rearranged pairs (e.g., ship-chilly), 'Inter' rearranged pairs (e.g., ship-pane), and 'Old-New' pairs (e.g., ship-bulb). Event-related potentials (ERPs) were recorded for each class of test item. Similar proportions of 'yes' responses to Old and Intra rearranged pairs were observed, but the false alarm rate for Inter rearranged pairs was markedly lower. ERPs elicited by correctly classified Old pairs and false alarms to Intra rearranged pairs were indistinguishable and exhibited 'left parietal' and 'right frontal' effects, characteristic of recollection-based recognition. ERPs for false alarms to Inter rearranged pairs did not show these effects. It is concluded that source judgment errors for confusable source-item pairings are mediated by the same processes that support correct source judgments. Such errors reflect recollection of gist information which is nondiagnostic with respect to source.

Analysis of Variance↗

PubMed2004

Probability effects on the neural correlates of retrieval success: an fMRI study.

Event-related fMRI was employed to investigate the influence of the relative probability of old and new test items on the neural correlates of recognition memory. Twelve subjects undertook three study-test cycles, each consisting of an identical study phase in which a series of words was encoded in an incidental task, followed by a test phase in which yes/no recognition judgments were made to a mixture of studied (old) and unstudied (new) words. The ratio of old to new words differed in each test phase, and was either 25:75, 50:50, or 75:25. In lateral inferior and medial parietal cortex, and the posterior cingulate, greater activity was elicited by correctly classified old than new items independently of old:new ratio. By contrast, in other regions, including anterior, dorsolateral, and ventrolateral prefrontal cortex, differences in the activity elicited by old and new items varied according to old:new ratio, demonstrating in some cases a complete crossover interaction. The results suggest that differential activity elicited by old and new test items is likely to support successful recognition in only a subset of the regions identified in previous studies as exhibiting such differences. In other regions, most notably prefrontal cortex, differences in the activity elicited by old and new items appear to reflect processes that are contingent upon, rather than in support of, successful recognition.

Adolescent↗