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PUNCHED CARD PROGRAMMING AND RECORDING TECHNIQUES EMPLOYED IN THE AUTOMATION OF THE WATA.

A system of punched card programming and recording of primate discrimination learning experiments is described to illustrate the application of these techniques to the automation of the discrete trial learning situation, allowing for punched card recording and subsequent digital computer data analyses without intermediate data reduction. The system was also designed to control several potential sources of variability inherent in the conventional WGTA testing mode, including 1) the subject's (S's) motivation to test on each trial, 2) S's attention to the stimuli, and 3) the rate of trial presentation so as to be more under S's control. Reliable techniques of punched card programming and recording are detailed.

Animals↗

Acetylcholine modulates averaged sensory evoked responses and perforant path evoked field potentials in the rat dentate gyrus.

The effect of localized application of acetylcholine (ACh) on well characterized components of sensory evoked and electrically induced potentials in the dentate gyrus was investigated in rats while performing a tone discrimination task. Local pressure application of ACh to the granule cell layer of the dentate gyrus through the recording pipette increased the amplitude of perforant path evoked population spikes without changing the amplitude of the field EPSP. When the pipette was relocated to the outer molecular layer of the dentate gyrus (OM), ACh application decreased the amplitude of the perforant path field EPSP. Two major components of the averaged auditory evoked potential (AEP) recorded during criterion performance of the discrimination task were significantly changed by dendritic application of ACh. The N1 component of the OM AEP which has been shown to reflect perforant path synaptic activity decreased in amplitude while the N2 component which represents activity from septal connections, was significantly increased. These effects were not due to the pressure ejection procedure nor drug related changes in behavioral performance of the task. The results suggest that ACh may act to differentially modulate the synaptic excitability of dentate granule cells, allowing them to acquire responses to sensory stimulation during the establishment and maintenance of discrimination learning.

Acetylcholine↗

Environmental enrichment, neocortical ectopias, and behavior in the autoimmune NZB mouse.

New Zealand Black (NZB) mice have severe autoimmune disease and approximately 40% have cortical ectopias in layer I of sensorimotor cortex. Because the ectopias are similar to those found in dyslexics, NZB mice have been used as an animal model for developmental learning disorders. In addition, these mice have been used as a model of learning deficits associated with autoimmune disease. To determine whether early intervention would affect learning processes in NZB mice, they were reared after weaning in standard cages or enriched environments. They were given a battery of behavioral tests to measure learning, laterality, and activity, after which they were sacrificed and their brains examined for cortical ectopias. The tests sorted into two behavioral sets. Ectopia-associated behaviors included black-white discrimination learning and the Morris spatial maze. As a group, the mice performed well on these tasks. Ectopic mice had poorer performance than non-ectopics on these measures, and environmental enrichment countered the effects of the ectopias. Autoimmune-associated behavior involved two-way avoidance learning in a shuttlebox. Mice were uniformely poor on this task, ectopias did not affect behavior, and environmental enrichment was without benefit. Evidence from this and other studies shows that poor shuttlebox performance is related to the presence of autoimmune disease. Thus, autoimmune disease and cortical ectopias each appear to affect a separate set of behavioral processes. Environmental enrichment is most effective for behavioral impairments mediated via cortical ectopias, but is much less effective, if at all, if autoimmunity is the primary mediator of the impairments.

Animals↗

Behavioral consequences of neonatal injury of the neocortex.

Several strains of autoimmune mice spontaneously develop molecular layer ectopias that are similar in appearance to those seen in humans and are caused by disturbances in neocortical neuronal migration. These mice also exhibit behavioral anomalies, some of which correlate with ectopias, others with the immunological disorder. In this study, we induced neocortical ectopias (via puncture wounds) and microgyria (via freezing lesions) in the neocortex of 1-day-old (newborn) mice without immune disorders in an attempt to further disentangle the effects of autoimmunity and of cortical malformation on behavior. In addition, we wished to compare the behavioral effects of small ectopias to larger microgyric lesions. DBA mice were assigned at birth to receive either a puncture wound or freezing lesion of either the left or right hemisphere. An independent group was subjected to sham surgery. In adulthood, these mice were given a battery of tests designed to measure lateralization and learning capacity. Lesioned mice (irrespective of hemisphere or type of damage) performed poorly when compared to sham-operated animals in discrimination learning, in a spatial Morris Maze Match-to-Sample task, and in a Lashley Type III maze. In shuttlebox avoidance conditioning, where immunological disorder has been shown to compromise behavioral performance in autoimmune mice, there was no difference between lesioned and sham animals. These results (1) support the dissociation between the effects of developmental neocortical anomalies and autoimmune disease on behavior (2) reveal similarities between spontaneous and induced neocortical malformations and (3) fail to support a difference in behavioral effects between ectopias and microgyria.

Animals↗

Arousal and learning, and noncontingency detection in one-year-old infants.

The relationship between arousal and learning was studied with 6 infants 1 yr. of age on a two-choice discrimination learning task. The behavioral indices of arousal (activity level) and learning (response rate and reinforcements) correlated significantly (r, .44). Affective state correlated positively with activity level (r, .47) during the learning phase of the experiment. Two control conditions were presented following the learning phase of the experiment. Noncontingent (response independent) reinforcement produced significantly faster response deceleration than nonreinforcement. Affective response did not differ between nonreinforcement and noncontingent reinforcement conditions.

Acoustic Stimulation↗

Context-specific interference on reversal learning of a stimulus-response habit.

Learning occurs in a particular place and time. In most learning situations, information about the training context is encoded along with the task demands and solution. However, the extent to which context contributes to the acquisition and expression of a particular learned response is unclear. In the present paper we examined two fundamental issues underlying the importance of context information and its role in expression of discrimination learning and reversal learning. Rats were trained on a stimulus-response (S-R) habit task designed for the eight-arm radial maze and after reaching a set criterion different context manipulations were performed. Results from Section 2.2.1 revealed that although rats detected a change in context, the learning was not context specific. Results from Section 2.2.2 showed that S-R reversal learning was enhanced when animals were reversed in a context that was different from the one used during original training. Animals that were reversed in a different context showed a renewal effect to the initial S-R when brought back to the original training context.

Animals↗

Attenuation of context-specific inhibition on reversal learning of a stimulus-response task in rats with neurotoxic hippocampal damage.

Rats with hippocampal or sham lesions were trained on a stimulus-response task developed for the 8-arm radial maze. After reaching a stringent learning criterion, different context manipulations were performed. In Experiment I, the different groups were transferred to an identical radial maze in a different room to determine the context specificity of the discrimination learning. Experiment I revealed that although rats with hippocampal lesions did not show a normal context detection effect, the expression of the discrimination was not context dependent for either the lesion or sham groups. In Experiment II, animals were trained to criterion on the discrimination task and then both groups were divided into sub-groups based on whether they would experience reversal training in the same or different context from original training. Experiment II indicated that animals with hippocampal lesions and shams reversed in a different context were significantly enhanced in reaching the learning criterion compared to either counterparts that were reversed in the same context. Reversal learning in rats with hippocampal lesions was faster than sham animals in the same context suggesting that the context-specific inhibition effect was hippocampal-based. After learning the reversal task, the groups of animals trained and reversed in different contexts were brought back into the original training context to test for competitive effects. Animals with hippocampal lesions that were reversed in the different context, did not show a competition between the most recently acquired discrimination and a context-specific association acquired during original training whereas sham animals in the same condition did. Taken together these results suggest that rats with hippocampal lesions do not acquire normal context-specific inhibition during discrimination learning.

Animals↗

Spatial relational memory in 9-month-old macaque monkeys.

This experiment assesses spatial and nonspatial relational memory in freely moving 9-mo-old and adult (11-13-yr-old) macaque monkeys (Macaca mulatta). We tested the use of proximal landmarks, two different objects placed at the center of an open-field arena, as conditional cues allowing monkeys to predict the location of food rewards hidden in one of two sets of three distinct locations. Monkeys were tested in two different conditions: (1) when local visual cues marked the two sets of potentially baited locations, so that monkeys could use both local and spatial information to discriminate these locations from never-baited locations; and (2) when no local visual cues marked the two sets of potentially baited locations, so that monkeys had to rely on a spatial relational representation of the environment to discriminate these locations. No 9-mo-old or adult monkey associated the presence of the proximal landmarks, at the center of the arena, with the presence of food in one set of three distinct locations. All monkeys, however, discriminated the potentially baited locations in the presence of local visual cues, thus providing evidence of visual discrimination learning. More importantly, all 9-mo-old monkeys tested discriminated the potentially baited locations in absence of the local visual cues, thus exhibiting evidence of spatial relational learning. These findings indicate that spatial memory processes characterized by a relational representation of the environment are present as early as 9 mo of age in macaque monkeys.

Age Factors↗

Stimulus function in simultaneous discrimination.

In discrimination learning, the negativity of the stimulus correlated with nonreinforcement (S-) declines after 100 training trials while the stimulus correlated with reinforcement (S+) is paradoxically more positive with lesser amounts of discrimination training. Training subjects on two simultaneous discrimination tasks revealed a within-subjects overlearning reversal effect, where a more-frequently presented discrimination problem was better learned in reversal than was a discrimination problem presented less frequently during training.

Animals↗

Quantitative analysis of risk sensitivity in honeybees (Apis mellifera) with variability in concentration and amount of reward.

Free-flying honeybees (Apis mellifera) were trained in a series of experiments designed to look for evidence of risk sensitivity in foraging for sucrose solution. The suitability of the choice method used was established in 3 preliminary experiments with differences in concentration, amount, and probability of reward. Of 5 subsequent experiments in which 2 alternatives provided the same mean concentration of sucrose solution with different variance, 3 showed risk indifference, and 2 showed risk aversion (preference for consistent reward). Of 2 final experiments in which the alternatives provided the same mean amount of sucrose solution with different variance, both showed risk aversion. Performance could be simulated quantitatively with a simple choice model developed by P. A. Couvillon and M. E. Bitterman (1991) to account for the results of a wide range of previous experiments on discriminative learning in honeybees.

Animals↗

Ontogenetic dissociation between habit learning and recognition memory in capuchin monkeys (Cebus apella).

The performance of young and adult capuchin monkeys (Cebus apella) on a Concurrent Discrimination Learning (CDL) test and a Delayed Non-Matching to Sample (DNMS) task were investigated. Results indicate that all subjects were able to learn the CDL test with 20-pairs simultaneously and retain this stimulus/reward association within 24-h interval. In contrast, young subjects did not perform the DNMS task with the same proficiency as adults. While adults' scores were above chance across all memory test delays, the young capuchin monkeys performed the test by chance level. These results support the hypothesis that these two tasks require different cognitive processes mediated by two independent neural systems with a differentiated ontogenetic development. Moreover, they provide evidence that this dissociation occurs not only in humans and Old World monkeys but also in the New World capuchin monkeys indicating that this species can be a valuable alternative model for investigations of the neurobiological basis of memory.

Age Factors↗

Interhemispheric communication of visual learning: a developmental study in 3-6-month old infants.

Infants aged 12-26 weeks (3-6 months) were presented with a visual category discrimination learning task (normal vs scrambled faces) in the right or left visual field. Once they had reached the learning criterion, they were tested for transfer to the untrained visual field in order to establish whether visual experience can be communicated to the opposite hemisphere. Transfer of learning was found to occur in infants aged 19-26 weeks but not in younger ones. Neural pathways possibly forming the channels for this communication are discussed.

Aging↗

Age-related changes in visual acuity, learning and memory in C57BL/6J and DBA/2J mice.

The DBA/2J mouse is a model of age-related pigmentary glaucoma in humans. Visual detection, pattern discrimination and visual acuity were evaluated in DBA/2J, C57BL/6J, B6.mpc1d (a C57 congenic strain) and D2.mpc1b (a D2 congenic strain) mice at 6, 12, 18 and 24 months of age. Mice were also tested in the Morris Water Maze and olfactory discrimination learning task. At 6 months, DBA/2J and D2.mpc1b mice outperformed C57BL/6J and B6.mpc1d mice in the visual detection task and there were no strain differences in performance on the water maze. At 12, 18 and 24 months, C57BL/6J and B6.mpc1d mice outperformed DBA/2J and D2.mpc1b mice in the vision tasks and in the water maze. Strains did not differ in the olfactory learning task. Therefore, loss of visual function occurs between 6 and 12 months of age in DBA/2J mice. Strain differences in visual task performance accounted for a significant proportion of the variance in measures of learning and memory in the water maze at 12, 18 and 24 months of age.

Aging↗

Visual learning for an auditory secondary reinforcer by macaques is intact after uncinate fascicle section: indirect evidence for the involvement of the corpus striatum.

Three cynomolgus monkeys (Macaca fascicularis) were trained preoperatively in visual discrimination learning for an auditory secondary reinforcer. Each new discrimination problem was solved on the basis of the secondary reinforcer, and primary reinforcement (food reward) was given only after a new problem had been solved. The animals learned 50 new problems in each daily session and it was therefore possible to assess accurately their average rate of learning new discrimination problems in this procedure. After the learning rate had stabilized preoperatively the animals were operated upon to transect the uncinate fascicle, the cortico-cortical pathway from visual association cortex in the temporal lobe to prefrontal cortex. The animals' learning rate was unchanged after uncinate fascicle section. A previous experiment has shown that visual learning for an auditory secondary reinforcer is unaffected by disconnection of visual association cortex from the amygdala and the fornix. Taken together, this negative evidence points strongly to the conclusion that visual learning for an auditory secondary reinforcer depends upon interaction of temporal lobe visual association cortex with the corpus striatum, since other possibilities have been excluded.

Acoustic Stimulation↗

An early and a late developing system for learning and retention in infant monkeys.

On the evidence that memory formation and habit formation represent two qualitatively different learning processes based on separate neural mechanisms, the functional development of these two processes was followed ontogenetically. Separate groups of rhesus monkeys of different ages were tested in delayed nonmatching-to-sample and 24-hr concurrent discrimination learning, considered to be measures of recognition memory and discrimination habit formation, respectively. The youngest group of infant monkeys failed to learn the nonmatching task until they were approximately 4 months old. With further maturation, learning ability on this task gradually improved, yet it did not reach adult levels of proficiency even at 1 year of age. Postlearning evaluation with long delays and lists confirmed this slow ontogenetic development of recognition memory to adult levels of function. By contrast, infant monkeys 3-4 months old learned to discriminate long lists of object-pairs about as quickly as adult monkeys despite the use of 24-hr intertrial intervals. This striking dissociation in the ability of infants on the two tasks closely resembles the dissociation first found in adult monkeys rendered amnesic by limbic lesions. The results suggest that whereas the nonlimbic habit system matures early in infancy, the limbic-dependent memory system develops only slowly.

Age Factors↗

The alpha(2A)-adrenergic agonist guanfacine improves visuomotor associative learning in monkeys.

Two monkeys were trained on two-problem visuomotor associations: if the cue was a circle pattern, move a handle to the left ('go-left'), and if it was a triangle pattern, move the handle to the right ('go-right'). These two visuomotor associations were unchanged throughout all the experiments and therefore were very familiar to the monkeys. For learning of new visuomotor associations, each monkey was presented with a new set of four novel patterns in each and every daily session, two of which instructed 'go-left' response and the other two 'go-right' response. Systemically administered guanfacine, a selective alpha(2A)-adrenergic agonist, improved the monkeys' learning ability: trials and errors to the learning criterion of 90% correct decreased significantly. The monkeys showed an enhanced capability of using at least three response strategies: win-stay on 'repeat trial', change-stay and change-shift on 'change trial'. The beneficial effect could be reversed by the coadministered idaxozan, an alpha(2)-adrenergic antagonist, which had no effect when administered alone. Similar treatment with guanfacine had no beneficial effect on visual discriminative learning, a task that involves the inferotemporal cortex. The present results indicate that stimulation by guanfacine of alpha(2A)-adrenoceptors improves visuomotor associative learning, probably through actions at alpha(2A)-adrenoceptors in the prefrontal cortex.

Adrenergic alpha-2 Receptor Agonists↗

Pavlovian conditioning of social-affiliative behavior in the Mongolian gerbil (Meriones unguiculatus).

Gerbils learned to approach a spatial-olfactory stimulus that signaled access to their pairmate. Experiments 1 and 3 used a discrimination procedure in which 1 conditioned stimulus (the CS+) was presented immediately before access to the pairmate and another (the CS-) was presented alone. Both male and female gerbils came to approach the CS+ sooner than the CS- and spent more time near the CS+ than the CS-. Discrimination learning was facilitated by making the CS+ and CS- spatially distinct (Experiment 3). Learning also was demonstrated in male gerbils, using a between-subjects design with a single CS. Pairing the CS with the opportunity for social interaction resulted in greater approach to the CS within 10 trials than presenting the CS and social opportunity in an unpaired fashion (Experiment 2). These findings demonstrate social-affiliative learning in the Mongolian gerbil. Similarities and differences between these findings and sexual conditioning effects in other species are discussed.

Animals↗

Conditioning of an interoceptive drug stimulus to different exteroceptive contexts.

Two experiments are described showing that a pentobarbital versus saline discrimination can be differentially conditioned to different external contexts. In experiment 1, rats differentiated two T-shaped mazes (one water maze and one electrified maze). In experiment 2, the external stimuli were the presence and absence of light during training in the electrified T-maze. These experiments constitute the first examples where a single drug versus nondrug discrimination has been conditioned to different external contexts requiring an opposite response pattern by the same animal. The data are in accordance with a stimulus interpretation of drug discrimination learning.

Animals↗