Marc W. Howard

Theta and Gamma Oscillations during Encoding Predict Subsequent Recall

Electrophysiological and hemodynamic measures of human brain activity have been shown to distinguish between episodes of encoding items that are later recalled versus those that are not recalled (Paller and Wagner, 2002). Using intracranial recordings from 793 widespread cortical and subcortical sites in 10 epileptic patients undergoing invasive monitoring, we compared oscillatory power at frequencies ranging from 2 to 64 Hz as participants studied lists of common nouns. Significant increases in oscillatory power during encoding predicted subsequent recall, with this effect predominantly in the 4-8 Hz (theta) and 28-64 Hz (gamma) frequency bands. Sites exhibiting increased theta activity during successful encoding were clustered in right temporal and frontal cortex, whereas those exhibiting increased gamma activity appeared bilaterally at widespread cortical locations. These findings implicate theta and gamma oscillatory activity, across a widespread network of cortical regions, in the formation of new episodic memories.

Contextual variability and serial position effects in free recall.

In immediate free recall, words recalled successively tend to come from nearby serial positions. M. J. Kahana (1996) documented this effect and showed that this tendency, which the authors refer to as the lag recency effect, is well described by a variant of the search of associative memory (SAM) model (J. G. W. Raaijmakers & R. M. Shiffrin, 1980, 1981). In 2 experiments, participants performed immediate, delayed, and continuous distractor free recall under conditions designed to minimize rehearsal. The lag recency effect, previously observed in immediate free recall, was also observed in delayed and continuous distractor free recall. Although two-store memory models, such as SAM, readily account for the end-of-list recency effect in immediate free recall, and its attenuation in delayed free recall, these models fail to account for the long-term recency effect. By means of analytic simulations, the authors show that both the end of list recency effect and the lag recency effect, across all distractor conditions, can be explained by a single-store model in which context, retrieved with each recalled item, serves as a cue for subsequent recalls.

Gradual Changes in Hippocampal Activity Support Remembering the Order of Events

The hippocampus is thought to contribute to episodic memory in part by binding stimuli to their spatiotemporal context. The present study examined how hippocampal neuronal populations encode spatial and temporal context as rats performed a task in which they were required to remember the order of trial-unique sequences of odors. The results suggest that a gradual change in the pattern of hippocampal activity served as a temporal context for odor-sampling events and was important for successful subsequent memory of the order of those odors.

A context-based theory of recency and contiguity in free recall

The authors present a new model of free recall on the basis of M. W. Howard and M. J. Kahanas temporal context model and M. Usher and J. L. McClellands leaky-accumulator decision model. In this model, contextual drift gives rise to both short-term and long-term recency effects, and contextual retrieval gives rise to short-term and long-term contiguity effects. Recall decisions are controlled by a race between competitive leaky accumulators. The model captures the dynamics of immediate, delayed, and continual distractor free recall, demonstrating that dissociations between short- and long-term recency can naturally arise from a model in which an internal contextual state is used as the sole cue for retrieval across time scales.

The Temporal Context Model in spatial navigation and relational learning: Toward a common explanation of medial temporal lobe function across domains

The medial temporal lobe (MTL) has been studied extensively at all levels of analysis, yet its function remains unclear. Theory regarding the cognitive function of the MTL has centered along 3 themes. Different authors have emphasized the role of the MTL in episodic recall, spatial navigation, or relational memory. Starting with the temporal context model (M. W. Howard & M. J. Kahana, 2002a), a distributed memory model that has been applied to benchmark data from episodic recall tasks, the authors propose that the entorhinal cortex supports a gradually changing representation of temporal context and the hippocampus proper enables retrieval of these contextual states. Simulation studies show this hypothesis explains the firing of place cells in the entorhinal cortex and the behavioral effects of hippocampal lesion in relational memory tasks. These results constitute a first step toward a unified computational theory of MTL function that integrates neurophysiological, neuropsychological, and cognitive findings.

Aging selectively impairs recollection in recognition memory for pictures: evidence from modeling and receiver operating characteristic curves.

Young and older adults were tested on recognition memory for pictures. The Yonelinas high threshold (YHT) model, a formal implementation of 2-process theory, fit the response distribution data of both young and older adults significantly better than a normal unequal variance signal-detection model. Consistent with this finding, nonlinear z-transformed receiver operating characteristic curves were obtained for both groups. Estimates of recollection from the YHT model were significantly higher for young than for older adults. This deficit was not a consequence of a general decline in memory; older adults showed comparable overall accuracy and in fact a nonsignificant increase in their familiarity scores. Implications of these results for theories of recognition memory and the mnemonic deficit associated with aging are discussed.

Shadows of the Past Temporal Retrieval Effects in Recognition Memory

We examine whether temporally defined associations play a role in item recognition. The role of these associations in recall tasks is well known; we demonstrate an important role in item recognition as well. In this study, subjects were significantly more likely to recognize a test item as having been previously experienced if the preceding test item was studied in a temporally proximal list position than if the preceding test item came from a more distant list position. Further analyses showed that this associative effect was almost entirely due to cases in which the preceding test item received a highest-confidence recognition judgment.

Ventral Hippocampal Neurons Are Shaped by Experience to Represent Behaviorally Relevant Contexts

Memories can be recalled at different levels of resolution, from a detailed rendition of specific events within a single experience to a broad generalization across multiple related experiences. Here we provide evidence that neural representations reflecting the specificity or generality of memories are differentially represented along the dorsoventral axis of the CA3 area of the rat hippocampus. In dorsal CA3, neurons rapidly associate the identity of events with specific locations whereas, in more ventrally located CA3 regions, neurons gradually accumulate information across extended training to form representations that generalize across related events within a spatial context and distinguish events across contexts.

Aging and contextual binding: Modeling recency and lag recency effects with the temporal context model.

Normal aging has been shown to spare recency effects in the initiation of free recall while disrupting temporally defined associations. The temporal context model (TCM) explains recency and temporally defined associations as consequences of a gradually changing context signal and recovery of those contextual states, respectively. Here we extend TCM to account for the dissociation between recency and temporally defined associations in younger and older adults. Modeling results suggested that the effect of aging was restricted to a decrement in the ability of items to recover the temporal contexts in which they were presented, a function that has been hypothesized to depend on the hippocampus.

Associative Retrieval Processes in Episodic Memory

Association and context constitute two of the central ideas in the history of episodic memory research. Following a brief discussion of the history of these ideas, we review data that demonstrate the complementary roles of temporal contiguity and semantic relatedness in determining the order in which subjects recall lists of items and the timing of their successive recalls. These analyses reveal that temporal contiguity effects normally studied on short timescales persist over very long timescales, a result that challenges traditional psychological and neuroscientific models of association. The form of the temporal contiguity effect is conserved across all of the major recall tasks and even appears in item recognition when subjects respond with high confidence. The near-universal form of the contiguity effect and its appearance at diverse timescales is shown to place tight constraints on the major theories of association.