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Dissociation of spatial reference memory, spatial working memory, and hippocampal mossy fiber distribution in two rat strains differing in emotionality.

Rats of the inbred strains DA/Han and BDE/Han were compared on two complex spatial learning tasks, a spatial reference memory task in a 16-unit multiple T-maze and a spatial working memory task in an eight-arm radial-maze. In addition, sizes of hippocampal mossy fiber terminal fields were measured. BDE rats showed marked superiority in multiple T-maze learning whereas DA rats outperformed BDE rats on the radial-maze task. DA rats had significantly larger intra- and infrapyramidal mossy fiber terminal fields (IIP-MF). This is consistent with findings from other studies suggesting that large IIP-MF are related to excellent spatial radial-maze learning, but it also indicates that size of IIP-MF is correlated with processing of a specific type of spatial information rather than with overall spatial abilities. BDE rats had more extended suprapyramidal mossy fiber projections (SP-MF) and a larger hilus. Rats of both strains differed in exploratory behaviour and emotionality: DA rats revealed little freezing and had a high rearing activity, whereas BDE rats showed frequent freezing and reared rarely. Results suggest that IIP-MF are involved with flexible expression of memory, updating environmental information and parallel processing whereas SP-MF might be linked to processing of familiar information. Presumably, emotional factors contribute to performance differences.

Animals↗

N-methyl-D-aspartate receptor antagonist MK-801 and spatial memory representation: working memory is impaired in an unfamiliar environment but not in a familiar environment.

Female Sprague-Dawley rats were injected with the noncompetitive N-methyl-D-aspartate (NMDA) antagonist MK-801 or saline 30 min before daily testing in spatial working memory (WM) and reference memory (RM) procedures in an 8-arm radial maze. MK-801 impaired RM and WM acquisition but not performance when rats were trained to criterion before drug administration. Neither a 2-hr nor a 4-hr delay between the first and last 2 correct WM choices impaired long-term WM. MK-801 impaired WM performance in trained rats only when rats were tested in a new environment. Thus, 2 mechanisms may be required for relational memory: an NMDA-dependent mechanism for acquiring long-term spatial representations and an NMDA-insensitive mechanism for operating on these stored representations.

Animals↗

How does the severity of a learning disability affect working memory performance?

Working memory performance was examined in children aged 11-12 years who had borderline, mild, and moderate learning disabilities. Comparisons with children of average abilities were used to determine whether those with more severe learning disabilities had greater impairments in working memory. Seven measures of working memory span were used to assess temporary phonological short-term storage (digit span, word span), temporary visuo-spatial short-term storage (pattern span, spatial span), and temporary short-term storage with additional processing, or central executive, demands (listening span. odd one out span, reverse digit span). Children with mild and moderate learning disabilities were impaired on all measures of working memory compared to children of average abilities. Children with borderline learning disabilities were just as good as children with average abilities on visuo-spatial and complex span tasks, but showed an impairment on phonological span tasks. Children with moderate learning disabilities were indistinguishable from children with mild learning disabilities on simple span tasks, but were significantly poorer than the mild group on the more demanding complex span tasks. For the group as a whole, working memory was strongly related to mental age.

Case-Control Studies↗

Working memory in another dimension: functional imaging of human olfactory working memory.

The majority of working memory research has been carried out within the visual and auditory modalities, leaving it unclear how other modalities would map onto currently proposed working memory models. In this study we examined the previously uninvestigated area of olfactory working memory. Our aim was to investigate if olfactory working memory would engage prefrontal regions known to be involved in working memory for other sensory modalities. Using positron emission tomography we measured cerebral blood flow changes in 12 volunteers during an olfactory working memory task and a comparison visual working memory task. Our findings indicate that both olfactory and face working memory engaged dorsolateral and ventrolateral frontal cortex when the task requirements were matched; a conjunction analysis indicated overlap in the distribution of activity in the two tasks. Similarities and differences in activity were noted in parietal lobe regions, with both tasks engaging inferior areas of 40/7, but only visual working memory showing increased activity within left superior parietal cortex. The findings support the idea that working memory processes engage frontal cortical areas independent of the modality of input, but do not rule out the possibility of modality-specific neural populations within dorsolateral or ventrolateral cortex.

Adult↗

Lidocaine reversible inactivation of the median raphe nucleus has no effect on reference memory but enhances working memory versions of the Morris water maze task.

Numerous studies in the past have dealt with the role of serotonergic system lesions in tasks aimed at measurement of cognitive behavior, but the literature concerning the role of serotonin in cognition remains controversial. Rats with electrolytic lesions of the median raphe nucleus (MRN) were found to display a profound impairment in both the acquisition and retention of spatial memory task. In this study, the lidocaine inactivation was employed to evaluate the involvement of the rat's median raphe nucleus in reference and working memory versions of the Morris water maze (MWM) task. Lidocaine (0.5 microl, 2%) was injected through a single cannula aimed at the MRN; control groups were treated in the same way with a 0.5 microl injection of saline. In Experiment 1, rats were trained in a reference memory version of the MWM with two blocks of four trials per day for three consecutive days, with intra-cerebral injection made 5 min before training. No significant difference was found. In Experiment 2, intra-cerebral injection was applied immediately after two blocks of four trials, and in Experiment 3, the drug was injected 5 min before retention test in rats that had received eight trials per day on three consecutive days. Again, no significant difference between control and treatment groups was found. These results indicate that MRN has no role in acquisition, consolidation and retrieval of spatial reference memory. In subsequent experiments, rats were trained in a working memory version of the MWM task to find a new target position in trial 1, and retrieval was tested 75 min later. MRN inactivation 5 min before (Experiment 4) and immediately after the acquisition trial (Experiment 5) enhanced spatial working memory. It is concluded that normal activity of the MRN has no role in formation and retrieval of reference memory, but it has an inhibitory role in working memory. Our results are confirmed with other studies suggesting that the serotonergic system has a different role in long-term and short-term memory. Interaction with other neurotransmitter systems like acetylcholine may be involved in this case.

Anesthetics, Local↗

The development of visuo-spatial working memory.

Children's performance on tests of visuo-spatial working memory improves with age, although relatively little is known about why this happens. One explanation concerns the development of the ability to recode visually presented information into phonological form. This process appears to be used from around 8 years of age and is a major contributor to tasks in which stimuli can be verbally labelled. However, evidence suggests that phonological recoding cannot account for all of the age-related change in performance on visuo-spatial working memory tasks. In this review, four other mechanisms (knowledge, processing strategies, processing speed, and attentional capacity) are considered in terms of their contribution to children's visuo-spatial working memory development.

Child↗

The effects of eye and limb movements on working memory.

Three experiments examined the role of eye and limb movements in the maintenance of information in spatial working memory. In Experiment 1, reflexive saccades interfered with memory span for spatial locations but did not interfere with memory span for letters. In Experiment 2, three different types of eye movements (reflexive saccades, pro-saccades, and anti-saccades) interfered with working memory to the same extent. In all three cases, spatial working memory was much more affected than verbal working memory. The results of these two experiments suggest that eye movements interfere with spatial working memory primarily by disrupting processes localised in the visuospatial sketchpad. In Experiment 3, limb movements performed while maintaining fixation produced as much interference with spatial working memory as reflexive saccades. These results suggest that the interference produced by eye movements is not the result of their visual consequences. Rather, all spatially directed movements appear to have similar effects on visuospatial working memory.

Extremities↗

Neuromodulatory control of interacting medial temporal lobe and neocortex in memory consolidation and working memory.

A model of the interacting medial temporal lobe and neocortical association areas is formulated and applied to memory consolidation and working memory. A simplified connectivity in terms of superassemblies and assemblies of neurons representing types and values, respectively, of object features is underlying the model. Realistic low-dimensional model neurons, developed in particular to take neuronal adaptation into account, are employed. Observed short- and long-term potentiation and depression of plastic synaptic couplings are incorporated. It is shown that memory consolidation by long-term potentiation, based on repeated activations of neocortical patterns, may be guided by neuromodulated dynamics of the medial temporal lobe via short-term couplings acting as pointers. Bifurcations, i.e. transitions between different modes of network dynamics, with respect to developing synaptic couplings are shown to depend on the adaptivity of excitatory neurons in the medial temporal lobe and thus to be under neuromodulatory control. At weak adaptivity, after an initial temporal segmentation of several objects accounting for the capacity of working memory to resolve several items, attention is selectively focused on a single object according to the model. At intermediate adaptivity, reactivations may persist and long-term synaptic couplings gradually develop. At strong adaptivity, the model predicts attention and memory consolidation to be subsequently terminated. The neuromodulatory control of the interacting medial temporal lobe and neocortical system via the adaptivity of excitatory neurons may account for several observations on the influence of neuromodulators on various cognitive processes and brain disorders.

Animals↗

Phonological deterioration in adults with an acquired severe hearing impairment: a deterioration in long-term memory or working memory?

The purpose of this study was to examine the phonological processing skills in individuals with an acquired severe hearing impairment. The subjects were tested on cognitive tasks that vary in their demands on phonological processing. The severely hearing impaired individuals performed at a significantly lower level on the word-word rhyme judgements task, but performed on a par with the control group on the picture-word rhyme judgements task and the lexical decision-making task. The results indicate that the phonological processing skills in individuals who have acquired a severe hearing loss in adult life deteriorate. The results are discussed with respect to theoretical and clinical implications.

Adult↗

Working memory and inferences: evidence from eye fixations during reading.

Eye fixations during reading were monitored to examine the relationship between individual differences in working memory capacity-as assessed by the reading span task-and inferences about predictable events. Context sentences predicting likely events, or non-predicting control sentences, were presented. They were followed by continuation sentences in which a target word represented an event to be inferred (inferential word) or an unlikely event (non-predictable word). A main effect of reading span showed that high working memory capacity was related to shorter gaze durations across sentence regions. More specific findings involved an interaction between context, target, and reading span on late processing measures and regions. Thus, for high- but not for low-span readers, the predicting condition, relative to the control condition, facilitated reanalysis of the continuation sentence that represented the inference concept. This effect was revealed by a reduction in regression-path reading time in the last region of the sentence, involving less time reading that region and fewer regressions from it. These results indicate that working memory facilitates elaborative inferences during reading, but that this occurs at late text-integration processes, rather than at early lexical-access processes.

Cognition↗

Word length and phonological similarity effects in simple, complex, and delayed serial recall tasks: implications for working memory.

Some current models of working memory argue that a passive short-term store is not involved in more dynamic working memory tasks. Other models argue that standard short-term memory and working memory tasks rely on common storage facilities. We examine these issues by exploring two signature effects of passive short-term storage in simple span, complex span, and Brown-Peterson tasks. The finding that all three tasks show word length and phonological similarity effects suggests that common processes or storage mechanisms are involved in all tasks. The implications for models of working memory are discussed.

Humans↗

Temporal dynamics of brain activation during a working memory task.

Working memory is responsible for the short-term storage and online manipulation of information necessary for higher cognitive functions, such as language, planning and problem-solving. Traditionally, working memory has been divided into two types of processes: executive control (governing the encoding manipulation and retrieval of information in working memory) and active maintenance (keeping information available 'online'). It has also been proposed that these two types of processes may be subserved by distinct cortical structures, with the prefrontal cortex housing the executive control processes, and more posterior regions housing the content-specific buffers (for example verbal versus visuospatial) responsible for active maintenance. However, studies in non-human primates suggest that dorsolateral regions of the prefrontal cortex may also be involved in active maintenance. We have used functional magnetic resonance imaging to examine brain activation in human subjects during performance of a working memory task. We used the temporal resolution of this technique to examine the dynamics of regional activation, and to show that prefrontal cortex along with parietal cortex appears to play a role in active maintenance.

Adult↗

Working memory in spelling: evidence from backward typing.

Theories of spelling (Margolin, 1984; Nolan & Caramazza, 1983) propose a working memory system for storing order and identity information of letters during the spelling process. Capacity limitations related to the use of such a graphemic buffer were explored. Participants had to type words backwards. Longer pauses between key presses were assumed to signal points at which graphemic buffer contents were refreshed. Five- and six-letter words were divided by a major pause into chunks of two and three letters, partly coinciding with syllables. Articulatory suppression had no effect on performance. Increasing the length of the stimuli to seven to eight letters resulted in major pauses occurring at syllable boundaries, and performance becoming vulnerable to articulatory suppression but not foot tapping. Forward typing resulted in a similar pause pattern. The results suggest that chunks of approximately three letters can be handled at any one time. For short words the task seems to rely on non-phonological modes of coding, whereas longer words appear to require the use of a phonological code, possibly for keeping track of progress through the word.

Humans↗

Organization of working memory within the human prefrontal cortex: a PET study of self-ordered object working memory.

The prefrontal cortex plays a critical role in working memory, the active maintenance of information for brief periods of time for guiding future motor and cognitive processes. Two competing models have emerged to account for the growing human and non-human primate literature examining the functional neuroanatomy of working memory. One theory holds that the lateral frontal cortex plays a domain-specific role in working memory with the dorsolateral and ventrolateral cortical regions supporting working memory for spatial and non-spatial material, respectively. Alternatively, the lateral frontal cortex may play a process-specific role with the more dorsal regions becoming recruited whenever active manipulation or monitoring of information in working memory becomes necessary. Many working memory tasks do not allow for direct tests of these competing models. The present study used a novel self-ordered working memory task and positron emission tomography to identify whether dorsal or ventral lateral cortical areas are recruited during a working memory task that required extensive monitoring of non-spatial information held within working memory. We observed increased blood flow in the right dorsolateral, but not ventrolateral, prefrontal cortex. Increases in blood flow in the dorsolateral region correlated strongly with task performance. Thus, the results support the process-specific hypothesis.

Adult↗

The hippocampus: a "working memory" structure? The effect of hippocampal sclerosis on working memory.

Patients with medically intractable epilepsy and either hippocampal sclerosis or frontal lobe lesions were compared with healthy controls, to investigate a possible neuroanatomical correlate of a component of working memory: the central executive. Patients were tested on a short-term memory task which comprised visuo-spatial and verbal components, in single and concurrent trials. Differences were found between the patient groups for dual-task capacity, despite being equated on single-task trials. Patients with frontal lobe damage were the most affected by the demands of attention division. The results of this study do not support the thesis of a hippocampal role in the working memory component examined, but point to a frontal lobe focus for this janusian cognitive function. An unexpected finding of an increment in performance over the trials of visuo-spatial assessment, in patients with hippocampal sclerosis, is presented.

Analysis of Variance↗

Neurocomputational models of working memory.

During working memory tasks, the firing rates of single neurons recorded in behaving monkeys remain elevated without external cues. Modeling studies have explored different mechanisms that could underlie this selective persistent activity, including recurrent excitation within cell assemblies, synfire chains and single-cell bistability. The models show how sustained activity can be stable in the presence of noise and distractors, how different synaptic and voltage-gated conductances contribute to persistent activity, how neuromodulation could influence its robustness, how completely novel items could be maintained, and how continuous attractor states might be achieved. More work is needed to address the full repertoire of neural dynamics observed during working memory tasks.

Action Potentials↗

Field dependence-independence from a working memory perspective: a dual-task investigation of the Hidden Figures test.

Field dependence-independence (FDI) is a construct intensively investigated within cognitive style research, but its cognitive underpinnings are not clearly specified. We propose that performance on FDI tasks primarily reflects the operations of the visuospatial and executive components of working memory. We tested this hypothesis in a dual-task experiment with a commonly used measure of FDI, the Hidden Figures Test. The results showed that performance on this test was impaired by concurrent performance of secondary tasks that primarily tap the visuospatial component (spatial tapping) and the executive component (2-back and random number generation), but was almost unaffected by other secondary tasks (simple tapping and articulatory suppression). Moreover, an analysis of secondary task performance ruled out the possibility of strategic trade-offs and revealed an intriguing dissociation for two different sets of "randomness" indices for the random number generation task. These results support the hypothesised mapping between FDI and working memory components and suggest that the dual-task paradigm can provide a useful way to bring underspecified constructs like FDI into closer alignment with theoretical ideas developed within cognitive psychology.

Field Dependence-Independence↗

Temporal interval production and processing in working memory.

Short-term memory or working memory has been proposed as a cognitive structure contributing to time estimation. Thus, in a previous experiment, retrieving a stored item during a temporal-interval production lengthened the interval in proportion to the number of items in the memory set. In the present study, this issue was analyzed further by testing whether the proportional lengthening is induced by the load itself (i.e., the number of items) or by comparing the probe with memorized items. In a first experiment, a memory set was maintained during a temporal production, and the comparison of the probe with memorized items was postponed until the end of time production. Varying the number of items in the memory set had no effect on temporal intervals produced during its retention, suggesting that mental comparison was the source of the lengthening of time intervals. In succeeding experiments, tasks requiring processing in working memory but involving no memory load were combined with temporal production. In Experiment 2, increasing the number of syllables in a rhyme-judgment task proportionally lengthened temporal intervals that were produced simultaneously. In Experiment 3, increasing the amount of mental rotation in a task involving visuospatial processing also lengthened simultaneous temporal production. This interference between processing in working memory and time estimation suggests that working memory, defined as a work space for active processing of current information, contributes to time estimation.

Adult↗