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PubMed1995

Lymphocyte memory and affinity selection.

The persistence of antigen-specific immune memory appears to require the presence of antigen--suggesting that memory may be due to restimulation of "memory" lymphocytes by persisting antigen. Persistence of antigen, in a form capable of stimulating B cell proliferation, on long-lived, follicular dendritic cells of lymphoid tissue is well documented. Existence of an analogous mechanism for T cell memory maintenance is controversial but can not be ruled out. Here we examine the consequences of immune memory maintenance by antigen-specific lymphocyte restimulation, and estimate the duration of memory as a function of model parameters. We show that the competition for restimulation among memory cell populations results in the selection of the clone having the highest overall affinity for the retained antigen. Thus affinity selection, an important attribute of immunity, is a constitutive property of memory maintenance by antigen-specific restimulation. In the case of B cells, affinity selection is predicted to continue to increase antibody affinity even after somatic mutation stops, and thus may be an important component of affinity maturation. Finally, we discuss several other hypotheses proposed to explain immune memory, including T cell stimulation by cross-reactive antigens.

Animals↗

PubMedDate not supplied

Memory and the hippocampus in food-storing birds: a comparative approach.

Comparative studies provide a unique source of evidence for the role of the hippocampus in learning and memory. Within birds and mammals, the hippocampal volume of scatter-hoarding species that cache food in many different locations is enlarged, relative to the remainder of the telencephalon, when compared with than that of species which cache food in one larder, or do not cache at all. Do food-storing species show enhanced memory function in association with the volumetric enlargement of the hippocampus? Comparative studies within the parids (titmice and chickadees) and corvids (jays, nutcrackers and magpies), two families of birds which show natural variation in food-storing behavior, suggest that there may be two kinds of memory specialization associated with scatter-hoarding. First, in terms of spatial memory, several scatter-hoarding species have a more accurate and enduring spatial memory, and a preference to rely more heavily upon spatial cues, than that of closely related species which store less food, or none at all. Second, some scatter-hoarding parids and corvids are also more resistant to memory interference. While the most critical component about a cache site may be its spatial location, there is mounting evidence that food-storing birds remember additional information about the contents and status of cache sites. What is the underlying neural mechanism by which the hippocampus learns and remembers cache sites? The current mammalian dogma is that the neural mechanisms of learning and memory are achieved primarily by variations in synaptic number and efficacy. Recent work on the concomitant development of food-storing, memory and the avian hippocampus illustrates that the avian hippocampus may swell or shrivel by as much as 30% in response to presence or absence of food-storing experience. Memory for food caches triggers a dramatic increase in the total number of number of neurons within the avian hippocampus by altering the rate at which these cells are born and die.

Animals↗

PubMed2002

Direct and indirect measures of verbal relational memory following anterior temporal lobectomy.

Medial temporal lobe (MTL) structures are implicated in forming conjunctions between events in order to form enduring relational memories; these memories are not evident using direct measures with varieties of amnesic subjects. Extratemporal brain structures are thought to be responsible for preserved memories, which are sometimes detectable using indirect measures. The present study tests this theory of multiple memory systems by examining whether preserved learning can be demonstrated for relational material in MTL-disordered subjects using an indirect measure which minimises conscious mediation of performance. The subjects had undergone anterior temporal lobectomy for relief of temporal lobe epilepsy: left-sided (LATL) cases had a mild verbal amnesia and right-sided (RATL) cases had better verbal memory, forming a comparison group. A direct measure of verbal relational memory was provided by successive trials of cued recall in a specially-constructed paired associate learning task with arbitrarily paired words; pairs consisted of either concrete or abstract words. LATL subjects performed worse than RATL subjects, and particularly so with abstract words. Following direct testing, memory for the pairings was measured indirectly using a masked recognition priming technique. RATL subjects showed savings in RT, demonstrating that masked priming can reveal evidence of the formation of conjunctions. Critically, LATL subjects showed no evidence of preserved learning with priming. Thus when MTL structures are damaged, relational memory appears to be affected without exception, consistent with the tenets of multiple memory systems theory.

Adult↗

PubMed2006

Upper alpha ERD and absolute power: their meaning for memory performance.

A variety of studies have shown that EEG alpha activity in the upper frequency range is associated with different types of cognitive processes, memory performance, perceptual performance and intelligence, but in strikingly different ways. For semantic memory performance we have found that resting or reference power is positively associated with performance, whereas during actual processing of the task, small power--reflected by a large extent of event-related desynchronization (ERD)--is related to good performance. We also have shown that the induction of large alpha reference power by neurofeedback training or repetitive transcranial magnetic stimulation (rTMS) at individual alpha frequency mimicked exactly the situation which is typical for good memory performance under normal situations: increased alpha reference power is associated with large ERD and good performance. Recent studies have demonstrated that this relationship holds true only for memory and not perceptual tasks that require the identification of simple visual stimuli under difficult conditions. In contrast to good memory performance, good perceptual performance is related to small pre-stimulus alpha power and a small ERD. We interpret this finding in terms of cortical inhibition vs. activation preceding task performance by assuming that large rhythmic alpha activity reflects inhibition. We assume that small reference alpha enhances perceptual performance because the cortex is activated and prepared to process the stimulus, whereas memory performance is enhanced if the cortex is deactivated before a task is performed because in typical memory tasks selective processing can start only after the to-be-remembered item or cue is presented. We also suggest that conflicting results about alpha ERD and the neural efficiency hypothesis (which assumes that highly intelligent exhibit a small ERD) can also be interpreted in terms of inhibition. Only if an intelligence test actually requires the activation of (semantic) memory, a large (because task specific) ERD can be observed. If other processing systems are required, the semantic memory system may even become suppressed, which is reflected by alpha event-related synchronization (ERS) or at least a largely decreased ERD.

Alpha Rhythm↗

PubMed1999

Source memory in major depression.

BACKGROUND: Research demonstrating episodic memory deficits in clinical depression has dealt with item memory exclusively. The present research sought to determine whether memory for source is differentially affected by depression. METHODS: Patients with major depression and normal control subjects were examined in item memory and two types of source memory, requiring discriminations between (e.g. something that the subject said and something another person said) and within (e.g. something that one person said and something another person said) classes. RESULTS: Depression-related deficits in item memory were exacerbated in source memory. However, deficits in source memory in depressed patients were restricted to those conditions requiring within-class discriminations. CONCLUSION: The overall pattern of results may reflect that those symptoms of major depression that affect the individuals basic processing resources (e.g. concentration difficulties, lack of effort, loss of energy) results in an impairment of episodic memory, particularly when the demands of differentiating perceptual and cognitive information are high.

Adult↗

PubMed2000

Emotional discomfort and impairments in verbal memory in schizophrenia.

Research has demonstrated that impairments in verbal memory in schizophrenia are linked with psychosocial deficits. Less is known, however, about their relationship to clinical features of illness. This study explores the hypothesis that impairments in verbal memory, particularly forms of memory requiring deeper levels of encoding, are uniquely linked to symptoms of dysphoria or emotional discomfort. Accordingly, we examined the association between concurrent measures of symptoms and verbal memory for 84 subjects with schizophrenia. Measures of positive, negative, cognitive, excitement and emotional discomfort symptoms were derived from factor scores of the Positive and Negative Syndrome Scale. Verbal memory was assessed using two tests requiring relatively superficial levels of encoding: The Hopkins Verbal Memory Test and the Digit Span subtest; and one test requiring deeper levels of encoding: the Logical Memory subtests I and II. As predicted, multiple regressions controlling for age, education and attention revealed that poorer performance on Logical Memory was strongly associated with greater levels of emotional discomfort (R(2)=0.22 and 0.25, respectively) while performance on the Hopkins test was related to cognitive symptoms scores (R(2)=0.08 and 0.09, respectively). Implications for the conceptualization of verbal memory deficits in schizophrenia are discussed.

Adult↗

PubMed2001

Multiple brain-memory systems: the whole does not equal the sum of its parts.

Most contemporary theories of memory are based on the assumption that memory can be divided into multiple psychological systems that are subserved by different neural substrates and that contribute to performance in a relatively independent manner. Although the study of individual memory systems has proved to be enormously useful, recent data increasingly point towards complex interactions between memory systems during performance of any given memory task. Three basic classes of interactions between different memory systems (competition, synergism and independence) are presented that appear to be congruent with the findings of many behavioral studies. Consideration of interactions among multiple memory systems will enhance our current understanding of memory by encouraging the view that memory systems are dynamic interactive units, rather than independent modules that act in isolation.

Animals↗

PubMed2001

New views on old memories: re-evaluating the role of the hippocampal complex.

Evidence of temporally graded retrograde amnesia (RA) following hippocampal damage has fuelled the long-standing belief that memory undergoes a consolidation process, whereby memories are progressively modified in neocortical regions until they are independent of the hippocampal (HPC) complex. Support for this position derives from both the animal and human RA literature, although the results are not consistent. Specifically, consolidation theory does not account for loss of episodic (detail) information in humans and context-dependent information in animals, which often extend back for much of the life span. We discuss an alternative approach, the Multiple Trace Theory, which suggests that the HPC complex contributes to the retrieval of recent and remote episodic and context-dependent memories. According to this view, such memory traces are represented as spatially distributed interactions between the HPC and neocortex that persist for as long as those memories exist. On the other hand, semantic, or context-free, memories can become independent of the HPC as consolidation theory predicts. In support of this view, we report recent accounts of relatively flat RA gradients in autobiographical and spatial detail loss in patients and animal models with extensive bilateral HPC lesions. By comparison, temporally graded RA was observed in tests of semantic and context-free memory. We also report neuroimaging studies in which hippocampal activity, elicited during recollection of autobiographical memories, did not distinguish recent from remote episodes. Our discussion suggests ways to reconcile discrepancies in the literature and guide predictions of the occurrence of flat versus temporally limited gradients of remote episodic and semantic memory loss following lesions to HPC.

Amnesia, Retrograde↗

PubMed2003

Facilitatory effect of the dopamine D4 receptor agonist PD168,077 on memory consolidation of an inhibitory avoidance learned response in C57BL/6J mice.

The still unknown contribution of the D4 receptors to memory consolidation was studied examining the memory effects of the dopamine D4 agonist PD168,077, the putative dopamine D4 antagonist L745,870, their mutual combination, and the combination of the D4 agonist with representative compounds acting as agonist or antagonist on the D1, D2 and the D3 receptors. Memory consolidation was assessed in C57BL/6J mice using the one-trial step-through inhibitory avoidance task, the compounds being injected immediately after training (foot-shock) and performance measured 24h later. PD168,077 (0.5-10mg/kg) dose-dependently improved memory performance and L745,870 (0.05-5mg/kg) at doses lower than 1mg/kg increased and at doses higher than 1mg/kg impaired memory performance. PD168,077 did not affect the paradoxical promnesic effect of low doses (0.1-0.5mg/kg) of L745,870, but antagonised the memory-impairing effect induced by 5mg/kg L745,870. The D1 antagonist SCH23390 (0.025-0.05 mg/kg) and the D2 antagonist eticlopride (0.01-0.05 mg/kg) antagonised the promnesic effects of PD168,077, which attenuated the decreasing effect on memory consolidation of both D1 and D2 antagonists. Accordingly, the D1 agonist SKF38393 (5-20mg/kg) and the D2 agonist quinelorane (0.1-1 mg/kg) both synergistically magnified the memory-improving effects of the D4 agonist. The dopamine D3 antagonist U99194A (2.5-10mg/kg) did not affect the promnesic effects induced by the D4 agonist, which nevertheless abolished the U99194A-induced promnesic effects. Additionally, the amnesic effects produced by the D3 agonist 7-OH-DPAT (0.01-1 microg/kg) was attenuated by PD168,077. These results suggest a potential role of dopamine D4 receptors in memory consolidation, which would be similar to that of the D1 and D2 receptors and probably opposite to that of the D3 receptors.

Analysis of Variance↗

PubMed1999

Memory dissociation: the approach to the study of retrieval processes.

The phenomenon of dissociated memory retrieval is observed when some influences (for example, pharmacological) on the brain result in specific changes of long-term memory. The purpose of present paper is to reveal possibilities of the phenomenon for study of long-term memory retrieval. Pharmacologically-induced dissociated states could be identified when the retrieval of responses learned before treatment is temporarily blocked by the drug influence, but the ability of the animals to learn new tasks is intact. Furthermore, memory traces that were formed in drugged state are not accessible for the retrieval in normal state and only the same drug treatment allows retrieving them. In the present work, dissociated learning of food-motivated tasks was carried out in Wistar rats with cholinesterase inhibitor physostigmine (0.5 mg/kg, intraperitonealy) or general anaesthetic sodium pentobarbital (15 mg/kg, intraperitonealy.). The retrieval of dissociated responses was studied under the influence of various doses of the same drugs. The results revealed the asymmetry of memory dissociation with physostigmine in contrast to pentobarbital-induced memory dissociation. Gradual access for the retrieval of dissociated memory traces after pharmacological modulation of cholinergic and GABA-ergic brain systems was shown. It was suggested an important role of hippocampus in memory dissociation, as a structure-performing match-mismatch operations between different retrieved memory traces.

Amnesia↗

PubMed2000

Concurrent modulation of anxiety and memory.

We have previously shown that the ventromedial prefrontal cortex (vmPFC) is involved in spontaneous working memory and anxiety-related behaviour in CD-1 mice. Specifically, pretrial microinjection of the kappa(1) agonist, U-69,593, in the infralimbic (IL) area of the vmPFC produced a robust anxiolytic behavioural profile in the elevated plus-maze and enhanced spontaneous working memory in the Y-maze. In the present study we sought to determine whether these effects were specific to IL kappa receptors. We hypothesized that microinjection of the kappa antagonist, norBNI, in the IL cortex would influence anxiety and spontaneous memory in an opposite direction to the effects produced by the kappa(1) agonist. In week 1, transfer-latency reference memory and anxiety were tested in the elevated plus-maze in two separate trials with an intertrial interval of 24 h. In week 2, spontaneous working memory was tested in the Y-maze followed immediately by defensive/withdrawal anxiety in the open field for one half of the animals in each group, and the other half was tested in reverse order. Pretreatment with one injection of vehicle, 1, 5 or 10 nmol/0.5 microl norBNI in the IL cortex dose-dependently reduced transfer-latencies and produced an anxiogenic behavioural profile in the first elevated plus-maze trial. Following a 24 h delay, transfer-latency reference memory was not influenced, but a robust anxiogenic behavioural profile was observed in the second no-injection anxiety trial in the elevated plus-maze relative to control animals. In week 2, the same groups of mice were again pretreated with one injection of the same doses of norBNI in the IL cortex and tested in the open field and Y-maze. NorBNI pretreatment was anxiogenic in the defensive/withdrawal anxiety test and disrupted spontaneous working memory regardless of testing order. The present results show the influence of kappa receptor modulation on anxiety induction and spontaneous working memory. These results also support the hypothesis that immediate memory processing may modulate the induction of anxiety-related behaviours.

Animals↗

PubMed1998

Effects of memory load on event-related patterns of 40-Hz EEG during cognitive and motor tasks.

The present study describes a phasic event-related synchronization (ERS) in the gamma band (36-44 Hz) induced by the onset of probe visual stimuli. The experiment consisted of two experimental tasks with high and low memory-load in which a geometrical figure (S1) was held in memory for comparison with a subsequent stimulus (S2). In each task a Go-NoGo paradigm was used with a Same-Different discrimination task. The aim of the study was to examine the influence on 40-Hz ERS of stimulus type ('Same-Different') and of response (Go-NoGo) task as moderated by activation or effort level (high/low memory-load). The investigation was carried out in 27 women. The EEG was recorded from FP1, FP2, F3, F4, P3, P4, O1 and O2 scalp sites referenced to linked earlobes. As a manipulation check for activation level, we recorded heart rate (HR) during the S1-S2 period. A main peak of activity was found around 160 ms with a maximum at occipital sites. The amplitude of this peak was higher in the high memory-load as compared to the low memory-load condition. This difference was manifested mainly at F3 and O2 scalp sites. A larger 40-Hz peak at F3 was also found in the Go compared to the NoGo condition. No 40-Hz ERS differences between Same and Different trials were observed. The HR was found sensitive to the stimulus type showing a greater HR deceleration response to S2 for Same trials, as compared to Different ones. In parallel with 40-Hz ERS response, the HR deceleration was more pronounced for the high memory-load as compared to the low memory-load condition. The results indicate that the 40-Hz ERS is dependent upon both memory-load and motor responding. The influence of memory load on cognitive (Same-Different) and motor response (Go-NoGo) variables is discussed.

Adult↗

PubMed2003

Age-related working memory impairment is correlated with increases in the L-type calcium channel protein alpha1D (Cav1.3) in area CA1 of the hippocampus and both are ameliorated by chronic nimodipine treatment.

The hippocampus is critical for spatial memory formation in rodents. Calcium currents through L-type voltage-sensitive calcium channels (L-VSCCs) are increased in CA1 neurons of the hippocampus of aged rats. We have recently shown that expression of the calcium conducting L-VSCC subunit alpha(1D) (Ca(v)1.3) is selectively increased in area CA1 of aged rats. We and others have speculated that excessive Ca(2+) influx through L-VSCC may be detrimental to memory formation. Therefore, we investigated the relationship between age-related working memory decline and alpha(1D) protein expression in the hippocampus. In addition, we studied the effects of chronic treatment with the L-VSCC antagonist nimodipine (NIM) on age-related working memory deficits and alpha(1D) expression in the hippocampus. Here we report that age-related increases in alpha(1D) expression in area CA1 correlate with working memory impairment in Fischer 344 rats. Furthermore, we demonstrate that chronic NIM treatment ameliorates age-related working memory deficits and reduces expression of alpha(1D) protein in the hippocampus. The present results suggest that L-VSCCs participate in processes underlying memory formation and that increases in L-VSCC protein and currents observed with aging may play a role in age-related memory decline. Furthermore, the amelioration in age-related memory decline produced by NIM treatment may be mediated, at least in part, by reductions in the abnormally high levels of alpha(1D) protein in the aged hippocampus. These findings may have implications for patients with Alzheimer's disease, who show increased L-VSCC protein expression in the hippocampus, and for patients receiving chronic treatment with L-VSCC antagonists.

Aging↗

PubMedDate not supplied

Memory performance in healthy elderly without Alzheimer's disease: effects of time and apolipoprotein-E.

Transgenic mice expressing human APOE-epsilon4 develop an age-dependent decline in memory without pathological features of Alzheimer's disease (AD). This implicates APOE in the maintenance of memory during normal senescence, but parallel human studies are limited because longitudinal investigations of memory usually do not exclude patients with AD or "questionable" AD (QD). The current study examined the effect of APOE on cognitive function over time in elderly without dementia. We hypothesized that, compared to other APOE alleles memory decline even in healthy elderly would be greater among those with an APOE-epsilon4. The results of neuropsychological tests, grouped into domains of memory, language and visuospatial/cognitive function by factor analysis, were examined at three intervals over a seven-year period in 563 healthy elderly without AD or QD using generalized estimating equations. Memory performance declined over time, while scores on the visuospatial/cognitive and language factors did not change. Increased age was associated with lower scores, and higher education with higher scores on all factors at each interval. No APOE allele was associated with performance on a specific cognitive factor at any interval, but the presence of an APOE-epsilon4 allele was associated with a more rapid decline in the memory factor over the follow-up period. The effect was most pronounced among individuals with less than 10 years of formal education. There was no similar time-dependent relationship between APOE-epsilon4 and the language or visuospatial/cognitive factors. Transgenic mice and elderly humans without AD or QD expressing APOE-epsilon4 show a decline in memory performance over time. These observations provide evidence for an APOE-specific effect on memory during senescence.

Aged↗

PubMed2001

Computational approaches to the architecture and operations of the prefrontal cortical circuit for working memory.

1. This article reviews recent progress in the computational studies towards the architecture and operations of the prefrontal cortical circuit, which are keys to understand the mechanisms of working memory processing. 2. The recurrent excitatory connections form closed-loop circuits, which contribute to the sustainment of delay-period activity. These connections subserve the cortical amplification of the activity. 3. Recent experimental studies (Wilson et al. 1994; Rao et al. 1999, 2000) suggested that at least two architectonically distinct types of intracortical inhibition, isodirectional and cross-directional inhibition, play significant roles in the formation of memory fields. 4. Computer simulations of a prefrontal cortical circuit model (Tanaka 1999, 2000a) showed that the isodirectional inhibition in the model regulated the amplitude of memory fields (i.e., the maximum firing rate) while the cross-directional inhibition contributed to the sharpening of the memory fields or the tuning curves. 5. The above characteristics enable the prefrontal cortical circuit to control memory fields, which would be necessary to general working memory processing. It would also be interesting to know whether different subtypes of the interneurons have distinct roles. 6. Another important issue is how neuromodulators contribute to working memory processing. Recent computer simulations by Durstewitz et al. (1999, 2000) showed that stronger dopamine action required stronger intervening input to destroy working memory, suggesting that dopamine contributes to the stabilization of working memory representation. 7. Further elucidation of these issues based on more detailed anatomical data of the cortical circuitry would make the architecture and operations of the prefrontal cortical circuit be more clearly described.

Animals↗

PubMed1999

Memories in context.

Context-dependent associative memories are models that allow the retrieval of different vectorial responses given a same vectorial stimulus, depending on the context presented to the memory. The contextualization is obtained by doing the Kronecker product between two vectorial entries to the associative memory: the key stimulus and the context. These memories are able to display a wide variety of behaviors that range from all the basic operations of the logical calculus (including fuzzy logics) to the selective extraction of features from complex vectorial patterns. In the present contribution, we show that a context-dependent memory matrix stores a large amount of possible virtual associative memories, that awaken in the presence of a context. We show how the vectorial context allows a memory matrix to be representable in terms of its singular-value decomposition. We describe a neural interpretation of the model in which the Kronecker product is performed on the same neurons that sustain the memory. We explored, with numerical experiments, the reliability of chains of contextualized associations. In some cases, random disconnection produces the emergence of oscillatory behaviors of the system. Our results show that associative chains retain their performances for relatively large dimensions. Finally, we analyze the properties of some modules of context-dependent autoassociative memories inserted in recursive nets: the perceptual autoorganization in the presence of ambiguous inputs (e.g. the disambiguation of the Necker's cube figure), the construction of intersection filters, and the feature extraction capabilities.

Memory↗

PubMed1999

Different forms of human odor memory: a developmental study.

Recognizing odors is an important biological function, both in the animal kingdom as well as for humans. It has been debated whether there exist different forms of human odor memory. For verbal memory, the concept of recollection and familiarity for conscious and unconscious recognition is widely accepted. Here we introduce a similar model for human odor memory. We use a combination of an odor naming and odor recognition memory task to estimate the relationship between depth of processing and retention of olfactory information. A developmental approach with children, young adults, middle aged adults and elderly subjects was chosen in order to study the influence of age. Our results indicate the existence of two separable forms of odor memory depending on whether the odors were correctly or incorrectly named during the naming task. These two forms of odor memory were differently represented across the human age range. Intact familiarity-based memory was found in all age groups, whereas memory based on recollection was impaired in the elderly and not yet fully developed in children. Our data show, for the first time, two different forms of human odor memory across the human life span.

Adolescent↗

PubMed2001

Frequency-specific effects of flicker on recognition memory.

The functions of the electroencephalographic rhythms are uncertain. Correlational evidence has linked 10-12Hz alpha rhythms to memory formation in healthy people. Moreover, loss of 10Hz alpha correlates with memory problems in Alzheimer's disease. Going beyond mere correlation, brain stimulation or peptides that alter electroencephalographic rhythms can modulate behaviour and enhance memory in rats. This latter finding fits with evidence that electroencephalogram-frequency stimulation can enhance long-term potentiation, the neural basis of memory. I aimed to test if manipulations of alpha-frequency electroencephalographic activity enhance human memory. Flicker provides an experimental means of modulating the human electroencephalogram: 9-12Hz flicker can entrain alpha-like activity. In rats, the frequency-specificity of brain stimulation's behavioural effects excludes the possibility that they result from non-specific (i.e. non-electroencephalographic) mechanisms. I tested if flicker manipulations would show analogous frequency-specific effects in man. In view of the above correlational data, I predicted that flicker at 10Hz (a frequency near to the peak power of endogenous alpha) would enhance human memory, but adjacent frequencies (8.7 and 11.7 Hz) would not. The results confirmed this prediction. This suggests that 10Hz electroencephalographic alpha subserves memory formation in man: 10Hz flicker enhances memory in healthy people and may have therapeutic potential in memory disorders.

Adult↗