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The effects of continuous versus partial reinforcement schedules on associative learning, memory and extinction in Lymnaea stagnalis.

A continuous schedule of reinforcement (CR) in an operant conditioning procedure results in the acquisition of associative learning and the formation of long-term memory. A 50 % partial reinforcement (PR) schedule does not result in learning. The sequence of PR-CR training has different and significant effects on memory retention and resistance to extinction. A CR/PR schedule results in a longer-lasting memory than a PR/CR schedule. Moreover, the memory produced by the CR/PR schedule is resistant to extinction training. In contrast, extinction occurs following the PR/CR schedule.

Animals↗

Associative learning impairments in patients with frontal lobe damage.

The performance of 18 frontal lobe lesion (FL) and 10 frontal lobe dementia (FLD) patients on an associative memory test was compared with the performance of their matched normal controls. The FL group was severely impaired on cued and free recall and was moderately impaired on a recognition condition. Left FL patients performed the poorest on the cued and free recall conditions. The FLD patients were moderately impaired on the free recall condition only but there was a subgroup of FLD patients with additional left temporal atrophy who appeared severely impaired on both cued and free recall. These findings indicate that both left frontal and temporal lobe damage can impair associative learning and that this impairment is more strikingly seen with free rather than cued recall.

Adult↗

Taste aversions to mother's milk: the age-related role of nursing in acquisition and expression of a learned association.

The development of taste aversion learning to novel cues contained in mother's milk was examined in rat pups. Pups receiving distinctive mild by experimenter-delivered oral infusions followed by toxicosis formed an aversion to the dam's diet. Robust aversions were learned as early as Day 10 and were retained for at least 11 days. When the same distinctive milk was obtained directly from a foster mother through nursing, however, only weanling-age pups (over 20 days) formed an aversion. X-ray analysis of nipple location in the mouths of suckling pups suggested that pups between the ages of 10 and 21 days receive milk at a similar tongue locus. Flavored milk was then delivered at specific time intervals in controlled quantities through tongue cannulas implanted at loci corresponding to the nipple position shown by the X-rays. Cannulated preweanling pups that were attached to a nipple during milk delivery failed to associate the taste cue with illness, whereas both preweanlings off the nipple and weanlings on the nipple acquired aversions to the taste cue in the milk. The evidence obtained in these experiments suggests that pups of all ages are incapable of expressing a taste aversion in a nursing situation and that preweanling pups in particular are also dificient in acquiring aversions within a suckling context. The inability of preweanling pups to acquire taste aversions in a nursing situation appears to result from a failure to associate taste cues with illness rather than a failure to detect taste cues obtained from a nipple.

Age Factors↗

Interaction of instructions with the recall strategy actually used in a paired-associates learning task.

Previous research suggests that interactive imagery produces memory performance superior to separation imagery, although the effects of instructions are less clear. Paired-associate learning was used to examine the effects of instructions (general memory, separation imagery, and interactive imagery) on recall, the frequency of using memory strategies (nonimaginal, separate image, and interactive image), and strategies' effectiveness. The numbers of correctly recalled items were fewer for subjects given general memory instructions than for subjects given separation or interactive imagery instructions which were not different from each other. Subjects reported using a variety of strategies. However, subjects given separation imagery instructions were more likely to report an interactive image than a separate image, and equally as likely to use interactive imagery as subjects given interactive imagery instructions. The present data suggest that subjects can effectively use a variety of memory strategies.

Adult↗

Single-cell neuronal model for associative learning.

Recently, a novel cellular mechanism, activity-dependent neuromodulation, was identified in sensory neurons mediating the gill and tail withdrawal reflexes in Aplysia. This mechanism may explain associative learning on a behavioral level. The present study was designed to mathematically model subcellular events that may underlie this mechanism and to examine the ability of the model to fit available empirical data. In this associative model, the reinforcing or unconditioned stimulus (US) leads to non-specific enhancement of transmitter release from sensory neurons by activating a cAMP cascade. Spike activity in sensory neurons, the conditioned stimulus (CS), transiently elevates intracellular Ca2+. The CS-triggered increases of intracellular Ca2+ "primes" the cyclase and amplifies the US-mediated cAMP synthesis. As a result of pairing specific amplification of cAMP levels, transmitter release is enhanced beyond that produced by unpaired stimuli or by application of the US alone. The model is capable of fitting empirical data on activity-dependent neuromodulation and predicts a characteristic interstimulus interval (ISI) curve. At the subcellular level, the model's ISI function is related to the time course of the buffering of intracellular Ca2+. The magnitude and duration of the pairing specific enhancement of transmitter release is related to the levels and time course of intracellular cAMP stimulation.

Action Potentials↗

Effects of sex and insulin/insulin-like growth factor-1 signaling on performance in an associative learning paradigm in Caenorhabditis elegans.

Learning is an adaptive change in behavior in response to environmental stimuli. In mammals, there is a distinct female bias to learn skills that is still unprecedented in other animal taxa. Here we have investigated the biological determinants of performance in an associative learning paradigm in the nematode Caenorhabditis elegans. Using an assay of chemotactic reactions associated with food deprivation, wild-type male worms show inferior learning ability relative to hermaphrodites. Sex-based learning difference is therefore an ancient evolutionary feature appearing even in relatively simple animals. C. elegans mutants with reduced insulin/IGF-1 signaling also exhibit a greatly reduced learning ability in this assay. In addition, hyperactivation of insulin/IGF-1 signaling through loss-of-function mutations in the PTEN phosphatase daf-18, a negative regulator of insulin/IGF-1 signaling, enhances learning ability beyond that of wild type. According to our epistasis analysis, the effect of DAF-2 on learning acts via phosphatidylinositol 3,4,5-trisphosphate (PIP(3)) production, but not the DAF-16 FOXO transcription factor. This implies that the signaling pathway from DAF-2 affecting this learning paradigm branches between PIP(3) production and DAF-16. However, learning capacity of nematodes is lowered by loss-of-function mutations in daf-16, suggesting involvement of noninsulin/IGF-1 signaling-dependent DAF-16 activation in learning. Potentially, sex and insulin/IGF-1 signaling affect performance in this learning assay via effects on the neurobiology of learning.

Animals↗

Effects on protein synthesis produced by pairing depolarization with serotonin, an analogue of associative learning in Aplysia.

A form of associative plasticity in Aplysia, activity-dependent neuromodulation, involves the convergence of neuronal activity and the effects of a modulatory transmitter. To investigate the role of protein synthesis in associative plasticity, we examined the effects of a biochemical analogue of activity-dependent neuromodulation on the level of incorporation of labeled amino acid into proteins. To mimic associative training, abdominal ganglia were exposed to paired treatments of a depolarizing agent, elevated potassium, and a modulatory transmitter, serotonin. The effects of elevated potassium and serotonin applied alone were also examined. At least two proteins (nos. 9 and 17) were affected in a nonadditive way by the paired procedure. Incorporation of label into protein 9 was increased by the paired procedure but was not affected by either elevated potassium or serotonin. Incorporation of label into protein 17 was significantly affected by elevated potassium or serotonin, but the effect of the paired procedure was significantly less than the summed effects of elevated potassium and serotonin applied alone. These results indicate that changes in protein synthesis may be important in the induction of associative plasticities. Amino acid sequences of two peptides derived from protein 9 were obtained. Then, a partial cDNA clone for protein 9 was obtained by performing PCR with degenerate primers corresponding to portions of the sequences of the two peptides. The sequence of protein 9 is related to sequences previously reported for a family of genes comprising the stringent starvation protein of Escherichia coli, auxin-induced proteins of plants, and glutathione S-transferases of a number of organisms.

Abdomen↗

Input and output changes of an identified neural pathway are correlated with associative learning in Hermissenda.

The effect of associative training on an identified putative motoneuron (MN1) in an identified visual input-output neural pathway was studied in Hermissenda crassicornis. The increased impulse frequency of the MN1 cell in response to a light step (MN1-LR) was found to be smaller up to 54 hr after associative training with paired light and rotation stimuli. No change was found in animals which received these training stimuli randomly. The MN1 activity in darkness, namely, baseline impulse frequency, spike amplitude, input resistance, and resting potential, did not change after training. The MN1-LR was positively correlated on retention days with the behavioral latency of the animal to enter the illuminated area of a light intensity gradient. However, the input resistance of the type B photoreceptor was inversely correlated with this behavioral latency as well as with MN1-LR. These findings suggest that the reduction of MN1-LR may be caused by the biophysical changes which have been found in the type B photoreceptor membrane. Furthermore, these findings support several other studies which indicate that specific biophysical changes in the type B photoreceptor membrane play a causal role in the observed behavioral modifications after associative training in Hermissenda.

Animals↗

Paired-associate learning in attention-deficit/hyperactivity disorder as a function of hyperactivity-impulsivity and oppositional defiant disorder.

A paired-associate learning (PAL) test was administered to 22 community volunteers without disruptive disorders and 197 children (7.5-13.5 years-old) presenting with the inattentive and combined subtypes of attention-deficit/hyperactivity disorder (ADHD) either in combination with or without oppositional defiant disorder (ODD). Participants were screened for learning disorders. In comparison to non-ADHD participants, children with ADHD achieved worse PAL and made errors rated as more acoustically and less semantically similar to the correct paired associates. These deficits were not related to hyperactivity-impulsivity or comorbid ODD. These results suggest that ADHD children are less competent at PAL and use less efficient learning strategies than their non-ADHD peers.

Adolescent↗

Paired associate learning in senile dementia of the Alzheimer type.

A study was conducted to examine the potential explicit learning ability in senile dementia of the Alzheimer type (SDAT) as a function of (1) the difficulty of the to-be-remembered material and (2) the severity of the dementia. The Associate Learning task from the Wechsler Memory Scale was administered to healthy older adults and individuals with questionable, mild, and moderate SDAT. Individuals with SDAT showed poorer learning performance across trials than healthy older adults. Individuals with questionable and mild SDAT did show evidence for learning across trials for highly related paired associates. For unrelated-paired associates, however, only individuals with questionable SDAT showed some evidence for increased learning across trials.

Alzheimer Disease↗

Fetal associative learning mediated through maternal alcohol intoxication.

BACKGROUND: The aim of the present study was to analyze whether alcohol as an unconditioned stimulus is capable of supporting associative learning in near-term fetuses. METHODS: In experiment 1, we determined pharmacokinetic profiles of alcohol and of an aromatic substance (cineole) in amniotic fluid and maternal blood during late gestation. The results obtained through gas chromatographic analysis allowed a second experiment in which we explicitly paired peak levels of cineole with peak levels of alcohol in amniotic fluid and blood, by intragastrically administering cineole and ethanol to the dams during gestational days 17 through 20 (paired condition). Control groups were dams given cineole 4 hr before commencement of an acute state of alcohol intoxication (long-delay group) or were only exposed to water administrations (water control group). The progeny were evaluated during postnatal day 16 in terms of behavioral responsiveness to intraorally infused solutions (cineole or alcohol presented in milk vehicle, or milk alone). RESULTS: Mouthing responsiveness to cineole was strongly affected by the nature of prenatal treatments. Pups in the paired prenatal condition mouthed significantly less than did long-delay and water controls. Physical and behavioral measures allowed us to reject the possibility that these effects were due to teratogenic effects of alcohol during late gestation. CONCLUSIONS: These results indicate that before birth, rat fetuses are capable of acquiring associative memories supported by the unconditioned properties of alcohol. This associative memory can be expressed during infancy through a significant reduction in mouth movements in the presence of the specific orosensory cue explicitly paired with alcohol interoceptive effects in utero.

Alcoholic Intoxication↗

Paired-associate learning in young and old adults as related to stimulus concreteness and presentation method.

This study investigated the learning of young and old adults as related to two variables, stimulus concreteness (low vs. high) and presentation method (recall vs. multiple choice vs. associate matching). The main findings were: (a) the elderly did not perform as well as the young adults, (b) for both age groups performance was better for the pairs with concrete stimuli than for the pairs with abstract stimuli, and (c) for both age groups performance was best with the associative matching method and poorest with the recall method. The paired-associate performance of the elderly is characterized by deficiencies in both response learning and associative learning.

Adult↗

Voltage-dependent calcium and potassium ion conductances: a contingency mechanism for an associative learning model.

Persistent light-induced depolarization results from Ca2+ influx across a photoreceptor membrane. The marked dependence on potential of this Ca2+ influx and a Ca+-dependent K+ efflux accounts for enhancement of the light-induced depolarization when light is paired with rotation. A positive feedback cycle between light-induced depolarization and synaptic depolarization due to stimulus pairing can explain long-lasting behavioral changes produced by associative training but not control paradigms. The sensitivity of this Ca2+ influx to intracellular levels of adenosine 3'-5'-monophosphate suggests biochemical steps for this model of associative learning.

Animals↗

Orbitofrontal cortex, associative learning, and expectancies.

Orbitofrontal cortex is characterized by its unique pattern of connections with subcortical areas, such as basolateral amygdala. Here we distinguish between the critical role of these areas in associative learning and the pivotal contribution of OFC to the manipulation of this information to control behavior. This contribution reflects the ability of OFC to signal the desirability of expected outcomes, which requires the integration of associative information with information concerning internal states and goals in representational memory.

Amygdala↗

Associative learning and representation. An EPS workshop for N. J. Mackintosh, 9 July 2002.

The papers published in this Special Issue are based upon presentations at a workshop on Associative Learning and Representation, which was sponsored by the Experimental Psychology Society and hosted by Emmanuel College, Cambridge. The workshop celebrated the contribution of Professor Nicholas Mackintosh to animal learning and conditioning in particular and experimental psychology in general in the year of his retirement from the Chair of Psychology at the University of Cambridge after 21 years in post. The date of the workshop, 9 July 2002, was particularly auspicious as it was the day of Professor Mackintoshs birth 67 years ago. Moreover, it is particularly fitting that this tribute is published in the Comparative and Physiological Psychology Section (B) of Quarterly Journal of Experimental Psychology, which he founded in 1981 during his editorship of the Journal between 1977 and 1984.

Animals↗

A stress-induced anxious state in male rats: corticotropin-releasing hormone induces persistent changes in associative learning and startle reactivity.

BACKGROUND: Exposure to intense inescapable stressors induces a persistent anxious state in rats. The anxious state is evident as increased sensory reactivity and enhanced associative learning. METHODS: We examine whether similar neurobehavioral changes are observed after intracerebroventricular (ICV) administration of corticotropin releasing hormone (CRH). Two behaviors were observed: acoustic startle responses (ASRs) and acquisition of the classically conditioned eyeblink response. Male Sprague-Dawley rats were administered ICV CRH either in a single dose (1.0 microg/rat) or in three doses each separated by 30 min. RESULTS: Exaggerated ASRs were evident 2 hours after either CRH treatment; however, only the rats given three injections exhibited a persistently exaggerated ASR apparent 24 hours after CRH treatment. Rats administered three injections of CRH also exhibited faster acquisition of the eyeblink conditioned response beginning 24 hours after treatment. Yet, we did not find evidence for a persistent activation of the HPA-axis response; three CRH injections did not lead to elevated basal plasma corticosterone levels the following morning. CONCLUSIONS: Repeated treatment with CRH over a 1.5-hour period models some of the behavioral changes observed after exposure to intense inescapable stressors.

Acoustic Stimulation↗

Prefrontal cortex and caudate nucleus in conditional associative learning: dissociated effects of selective brain lesions in rats.

Rats with lesions to prefrontal cortex (PFC) or caudate nucleus (CN) were compared on tests of conditional associative learning (CAL) that placed varying demands on conditional rule learning and working-with-memory operations that are essential for response selection. Damage to either structure impaired performance, but the respective deficits resulted from disruption of different processes. CN lesions produced a consistent learning deficit that was thought to reflect a basic impairment in forming stimulus-response (S-R) associations. The behavior of PFC rats was more variable and depended on task requirements. When S-R learning or response selection was relatively easy, the PFC was not critical. However, when either component was made more difficult, thus requiring the contribution of strategic processes, PFC damage produced profound impairments. In addition to clarifying the roles of the PFC and CN in CAL, the results provide further evidence that multiple brain regions participate in relatively simple behavioral tasks and that their respective contributions can be dissociated.

Animals↗

The p38 mitogen-activated protein kinase is involved in associative learning in rabbits.

This study examined the role of the mitogen-activated protein kinase (MAPK) family during acquisition of the rabbit's classically conditioned eye-blink response. Eye-blink conditioning produced a significant, bilateral activation of both extracellular signal-regulated protein kinases (ERKs) and p38 MAPK in the anterior cerebellar vermis. There was also a significant bilateral activation of ERKs in the dorsal hippocampus with no change in p38 MAPK. These changes were seen at 2 min after the last conditioning session, were maintained for at least 180 min, and occurred without any change in the protein expression of either ERKs or p38 MAPK. There were no changes in ERKs or p38 MAPK in frontal cortex, in cerebellar hemispheral lobule VI, or in a section of brainstem containing the inferior olive. Moreover, the stress-related protein kinase Jun N-terminal kinase (JNK), another subfamily of MAPKs, was not altered in any of the brain regions examined. Animals receiving explicitly unpaired presentations of a conditioned stimulus and an unconditioned stimulus did not acquire conditioned responses (CRs) and did not demonstrate any changes in ERKs, p38 MAPK, or JNK. The intraventricular injection of SB203580, a selective p38 MAPK inhibitor, significantly retarded CR acquisition and blocked the learning-related increases in p38 MAPK activity in the anterior vermis. PD98059, a selective MAPK kinase inhibitor, had a smaller and only marginally significant effect on CR acquisition, although it did block the learning-related increases in ERK activity in both the hippocampus and anterior vermis. These results indicate that p38 MAPK is activated during associative learning and may play a role in the transcriptional events that lead to memory consolidation.

Acoustic Stimulation↗