Behavioral and neurophysiological correlates of episodic coding,proactive interference,and list length effects in a running span verbal working memory task

Updating refers to (1) discarding items from, (2) repositioning items in, and (3) adding items to a running working memory span. Our behavioral and fMRI experiments varied three factors: trial length, proactive interference (PI), and group integrity. Group integrity reflected whether the grouping of items at the encoding stage was violated at discarding. Behavioral results were consistent with the idea that updating processes have a relatively short refractory period and may not fatigue, and they revealed that episodic information about group context is encoded automatically in working memory stimulus representations. The fMRI results did not show evidence that updating requirements in a task recruit executive control processes other than those supporting performance on nonupdating trials. They did reveal an item-accumulation effect, in which signal increased monotonically with the number of items presented during the trial, despite the insensitivity of behavioral measures to this factor. Behavioral and fMRI correlates of PI extended previous results and rejected an alternative explanation of PI effects in working memory.     

Integrating incremental learning and episodic memory models of the hippocampal region

By integrating previous computational models of corticohippocampal function, the authors develop and test a unified theory of the neural substrates of familiarity, recollection, and classical conditioning. This approach integrates models from 2 traditions of hippocampal modeling, those of episodic memory and incremental learning, by drawing on an earlier mathematical model of conditioning, SOP (A. Wagner, 1981). The model describes how a familiarity signal may arise from parahippocampal cortices, giving a novel explanation for the finding that the neural response to a stimulus in these regions decreases with increasing stimulus familiarity. Recollection is ascribed to the hippocampus proper. It is shown how the properties of episodic representations in the neocortex, parahippocampal gyrus, and hippocampus proper may explain phenomena in classical conditioning. The model reproduces the effects of hippocampal, septal, and broad hippocampal region lesions on contextual modulation of classical conditioning, blocking, learned irrelevance, and latent inhibition.     

Computational models of the hippocampal region: linking incremental learning and episodic memory

The hippocampal region, a group of brain structures important for learning and memory, has been the focus of a large number of computational models. These tend to fall into two groups: (1) models of the role of the hippocampal region in incremental learning, which focus on the development of new representations that are sensitive to stimulus regularities and environmental context; (2) models that focus on the role of the hippocampal region in the rapid storage and retrieval of episodic memories. Rather than being in conflict, it is becoming apparent that both approaches are partially correct and might reflect the different functions of substructures of the hippocampal region. Future computational models will help to elaborate how these different substructures interact.     

Neural mechanisms of interference control underlie the relationship between fluid intelligence and working memory span

Fluid intelligence (gF) and working memory (WM) span predict success in demanding cognitive situations. Recent studies show that much of the variance in gF and WM span is shared, suggesting common neural mechanisms. This study provides a direct investigation of the degree to which shared variance in gF and WM span can be explained by neural mechanisms of interference control. The authors measured performance and functional magnetic resonance imaging activity in 102 participants during the n-back WM task, focusing on the selective activation effects associated with high-interference lure trials. Brain activity on these trials was correlated with gF, WM span, and task performance in core brain regions linked to WM and executive control, including bilateral dorsolateral prefrontal cortex (middle frontal gyrus; BA9) and parietal cortex (inferior parietal cortex; BA 40/7). Interference-related performance and interference-related activity accounted for a significant proportion of the shared variance in gF and WM span. Path analyses indicate that interference control activity may affect gF through a common set of processes that also influence WM span. These results suggest that individual differences in interference-control mechanisms are important for understanding the relationship between gF and WM span.     

Intentional memory and online data collection: A test of the effects of animacy and threat on episodic memory

As research in the behavioural sciences increasingly relies on Amazon’s Mechanical Turk (MTurk) for recruiting participants, it is an empirical question whether an intentional memory experiment could be conducted online and whether performance of MTurk participants and college students would be comparable. College students and MTurk participants viewed threatening and nonthreatening items that were animate or inanimate and completed a recall test. Recent research indicates that human memory systems evolved to prioritise survival-relevant information, including animacy of items. The results of both studies indicated that participants recalled more animate than inanimate items and recalled more threatening items than nonthreatening items. The results indicate that it is possible to conduct memory experiments online and that results match patterns found in laboratory studies of the animacy effect (Nairne, VanArsdall, Pandeirada, Cogdill, & LeBreton, 2013), but that researchers should include safeguards to ensure the online participants are attending to and following directions.     

Mechanisms of aural encoding: VIII. Phonetic interference and context-sensitive coding in short-term memory

In previous experiments Ss were presented for ordered recall with sequences of five consonant phonemes paired with /a/ in which the middle three consonant phonemes shared the same manner of articulation (voiced, unvoiced, nasal), the same place of articulation (front, middle, back), or neither the same manner nor place of articulation (control sequences). Compared to performance in control sequences, the middle consonant phoneme was always more difficult to recall in manner of articulation sequences but not in place of articulation sequences. The results suggested that for these sequences consonant phonemes were not remembered in terms of their place of articulation. In the present experiment, sequences of consonant-vowel (CV) or vowel-consonant (VC) syllables were presented for recall in which each consonant phoneme was paired with a different vowel. When consonant phonemes in the different sequence types were presented for recall with different vowels, phonetic interference was observed for the middle consonant in place of articulation sequences as well as manner of articulation sequences, and the effect was observed in both CV and VC groups. It was suggested that vowels are encoded in short-term memory in terms of their place of articulation and that presenting consonant phonemes for recall with different vowels caused Ss to use this dimension to code consonant phonemes in short-term memory.     

Conjunctive representations in learning and memory: principles of cortical and hippocampal function

The authors present a theoretical framework for understanding the roles of the hippocampus and neocortex in learning and memory. This framework incorporates a theme found in many theories of hippocampal function: that the hippocampus is responsible for developing conjunctive representations binding together stimulus elements into a unitary representation that can later be recalled from partial input cues. This idea is contradicted by the fact that hippocampally lesioned rats can learn nonlinear discrimination problems that require conjunctive representations. The authors' framework accommodates this finding by establishing a principled division of labor, where the cortex is responsible for slow learning that integrates over multiple experiences to extract generalities whereas the hippocampus performs rapid learning of the arbitrary contents of individual experiences. This framework suggests that tasks involving rapid, incidental conjunctive learning are better tests of hippocampal function. The authors implement this framework in a computational neural network model and show that it can account for a wide range of data in animal learning.     

Low perceived control as a risk factor for episodic memory: the mediational role of anxiety and task interference

Low perceived control is considered a risk factor for poor cognitive functioning, but the mechanisms are unclear. The goal of the present study was to analyze anxiety and task interference as sequential mediators of the association between control beliefs and episodic memory. Cognitive-specific control beliefs were assessed prior to the lab session. State anxiety was assessed in the lab, followed by a word list recall task. The frequency of intrusive thoughts during the memory task was reported by the participants as a measure of task interference after the completion of the cognitive testing. The results for 152 participants from the ages of 22 to 84 years supported the predicted three-path mediation model. Lower levels of control beliefs were associated with higher state anxiety, which in turn affected episodic memory performance by increasing the likelihood of task interference, with age, sex, and verbal abilities as covariates. The implications of the results for developing interventions to improve memory performance are considered.     

Attentional distractor interference may be diminished by concurrent working memory load in normal participants and traumatic brain injury patients

A reduction in congruency effects under working memory (WM) load has been previously described using different attentional paradigms (e.g.,  and ). One hypothesis is that different types of WM load have different effects on attentional selection, depending on whether a specific memory load demands resources in common with target or distractor processing. In particular, if information in WM is related to the distractors in the selective attention task, there is a reduction in distraction (Kim et al., 2005). However, although previous results seem to point to a decrease in interference under high WM load conditions (Kim et al., 2005), the lack of a neutral baseline for the congruency effects makes it difficult to differentiate between a decrease in interference or in facilitation. In the present work we included neutral trials in the task introduced by Kim et al. (2005) and tested normal participants and traumatic brain injury patients. Results support a reduction in the processing of distractors under WM load, at least for incongruent trials in both groups. Theoretical as well as applied implications are discussed.     

Attentional distractor interference may be diminished by concurrent working memory load in normal participants and traumatic brain injury patients

A reduction in congruency effects under working memory (WM) load has been previously described using different attentional paradigms (e.g., Kim et al., 2005, Smilek et al., 2006). One hypothesis is that different types of WM load have different effects on attentional selection, depending on whether a specific memory load demands resources in common with target or distractor processing. In particular, if information in WM is related to the distractors in the selective attention task, there is a reduction in distraction (Kim et al., 2005). However, although previous results seem to point to a decrease in interference under high WM load conditions (Kim et al., 2005), the lack of a neutral baseline for the congruency effects makes it difficult to differentiate between a decrease in interference or in facilitation. In the present work we included neutral trials in the task introduced by Kim et al. (2005) and tested normal participants and traumatic brain injury patients. Results support a reduction in the processing of distractors under WM load, at least for incongruent trials in both groups. Theoretical as well as applied implications are discussed.     

Decision-making by experienced rugby referees: use of perceptual information and episodic memory

The expertise paradigm was used in two studies to examine decision-making by rugby referees. Videoclips were used to assess infraction detection and knowledge base, or sources of information used. In Study 1, referees of high and low experience were compared, and in Study 2 referees and players were compared. Analysis by signal detection used a framework to classify types of information. Study 1 yielded no differences in detection of infractions as a function of experience, however, referees of high experience used significantly more sources of information than the group with low experience across all categories of information. In Study 2, there were no significant differences between referees and players, with the exception that referees displayed greater use of episodic memory information in decision-making.     

Memory retrieval time and memory capacity of the CA3 network: role of gamma frequency oscillations

The existence of recurrent synaptic connections in CA3 led to the hypothesis that CA3 is an autoassociative network similar to the Hopfield networks studied by theorists. CA3 undergoes gamma frequency periodic inhibition that prevents a persistent attractor state. This argues against the analogy to Hopfield nets, in which an attractor state can be used for working memory. However, we show that such periodic inhibition allows one cycle of recurrent excitatory activity and that this is sufficient for memory retrieval (within milliseconds). Thus, gamma oscillations are compatible with a long-term autoassociative memory function for CA3. A second goal of our work was to evaluate previous methods for estimating the memory capacity (P) of CA3. We confirm the equation, P = c/a(2), where c is the probability that any two cells are recurrently connected and a is the fraction of cells representing a memory item. In applying this to CA3, we focus on CA3a, the subregion where recurrent connections are most numerous (c = 0.2) and approximate randomness. We estimate that a memory item is represented by approximately 225 of the 70,000 neurons in CA3a (a = 0.003) and that approximately 20,000 memory items can be stored. Our general conclusion is that the physiological and anatomical findings of CA3a are consistent with an autoassociative function. The nature of the information that is associated in CA3a is discussed. We also discuss how the autoassociative properties of CA3 and the heteroassociative properties of dentate synapses (linking sequential memories) form an integrated system for the storage and recall of item sequences. The recall process generates the phase precession in dentate, CA3, and entorhinal cortex.     

Differential interaction of platelet-activating factor and NMDA receptor function in hippocampal and dorsal striatal memory processes

The interaction between platelet activating factor (PAF) and NMDA receptor function in hippocampal and dorsal striatal memory processes was examined. In both a hidden and a visible platform water maze task, peripheral post-training injection of MK-801 (0.05 mg/kg) impaired memory. Post-training intrahippocampal infusions of PAF (1.0 microg/0.5 microl) enhanced memory in the hidden platform task, while intradorsal striatal infusion of PAF (1.0 microg/0.5 microl) enhanced memory in the visible platform task. The memory impairing effects of post-training injection of MK-801 was blocked by concurrent intrahippocampal infusion of PAF. In contrast, post-training injection of MK-801 blocked the memory enhancing effects of concurrent intradorsal striatal infusion of PAF. The results suggest that (1) the memory enhancing effects of intracerebral PAF infusion involve an interaction with NMDA receptor function, and (2) the nature of this interaction may represent a differential mechanism mediating the distinct roles of the hippocampus and dorsal striatum in cognitive memory and stimulus-response habit formation, respectively.     

Coding of stimuli by animals: Retrospection, prospection, episodic memory and future planning

When animals code stimuli for later retrieval they can either code them in terms of the stimulus presented (as a retrospective memory) or in terms of the response or outcome anticipated (as a prospective memory). Although retrospective memory is typically assumed (as in the form of a memory trace), evidence of prospective coding has been found when response intentions and outcomes are particularly salient. At a more abstract level is the question of whether animals are able figuratively to travel back in time to recover memories of past events (episodic memory) and forward in time to predict future events (future planning). Although what would constitute adequate evidence of episodic memory and future planning is controversial, preliminary evidence suggests that animals may be capable of both forms of subjective time travel.     

Working memory deficits in boys with attention deficit/hyperactivity disorder (ADHD): An examination of orthographic coding and episodic buffer processes

The episodic buffer component of working memory was examined in children with attention deficit/hyperactivity disorder (ADHD) and typically developing peers (TD). Thirty-two children (ADHD = 16, TD = 16) completed three versions of a phonological working memory task that varied with regard to stimulus presentation modality (auditory, visual, or dual auditory and visual), as well as a visuospatial task. Children with ADHD experienced the largest magnitude working memory deficits when phonological stimuli were presented via a unimodal, auditory format. Their performance improved during visual and dual modality conditions but remained significantly below the performance of children in the TD group. In contrast, the TD group did not exhibit performance differences between the auditory- and visual-phonological conditions but recalled significantly more stimuli during the dual-phonological condition. Furthermore, relative to TD children, children with ADHD recalled disproportionately fewer phonological stimuli as set sizes increased, regardless of presentation modality. Finally, an examination of working memory components indicated that the largest magnitude between-group difference was associated with the central executive. Collectively, these findings suggest that ADHD-related working memory deficits reflect a combination of impaired central executive and phonological storage/rehearsal processes, as well as an impaired ability to benefit from bound multimodal information processed by the episodic buffer.     

Facilitative and interference effects of facial expressions on familiarity as a function of repetition priming frequency

Facial expressions of familiar faces have been found to influence identification. In this study, we hypothesize that faces with emotional expression are encoded for both structural and variant information resulting in more robust identification. Eighty-eight participants were presented faces with repetition priming frequencies of 2, 5, 10, and 20 (learning stage) and then judged the faces in terms of familiarity (testing stage). Participants were randomized into one of the following conditions: The facial expression between learning and testing stage remained the same (F-F), faces with facial expression shown in the learning stage were neutralized in the testing stage (F-N), or faces with neutral expressions shown in the learning stage had emotional expressions in the testing stage (N-F). Results confirmed our hypothesis and implications are discussed.     

The effects of episodic and semantic memory on the identification of sentences masked by white noise

Two studies examine how episodic and semantic memory affect subjects’ abilities to repeat sentences masked by white noise. Subjects first hear a list of 70 sentences that are not masked. Subjects in the framework conditions are told prior to hearing the list that all sentences refer to a contextual framework concerning a deserted island. Subjects in the no-framework groups are not given this information. Subjects in the framework-after condition are given this information only after hearing the list of sentences. Subjects then perform a white-noise identification task. The results indicate that framework subjects are able to identify both old and new framework-related information better than other subjects. Subjects in the no-framework and framework-after conditions identify old information better than control subjects who do not participate in an acquisition phase. Emphasis is placed on the interdependence of episodic and semantic memory, including conditions leading to such interdependence.