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Schema theory (1975): retrospectives and prospectives.

A brief commentary is provided on the theoretical assumptions, scholarly impact and continuing influence of the schema theory of motor learning (Schmidt, 1975). The traditional contrasts of schema theory to the coordinative structure or dynamical systems framework are reemphasized, and limitations of the variability of practice experiments noted. A central problem for theories of motor learning is change over time, the basis on which learning is typically defined. Most theories including schema have, however, undervalued the importance of the time-dependent nature of change in deference to the almost exclusive study of the amount of some averaged change in behavioral outcome. The persistent and transitory change(s) in movement and outcome that are observed in action are reflections of multiple time scales of change in a dynamical system.

Humans↗

Comparative learning theory and its application in the training of horses.

Training can best be explained as a process that occurs through stimulus-response-reinforcement chains, whereby animals are conditioned to associate cues in their environment, with specific behavioural responses and their rewarding consequences. Research into learning in horses has concentrated on their powers of discrimination and on primary positive reinforcement schedules, where the correct response is paired with a desirable consequence such as food. In contrast, a number of other learning processes that are used in training have been widely studied in other species, but have received little scientific investigation in the horse. These include: negative reinforcement, where performance of the correct response is followed by removal of, or decrease in, intensity of a unpleasant stimulus; punishment, where an incorrect response is paired with an undesirable consequence, but without consistent prior warning; secondary conditioning, where a natural primary reinforcer such as food is closely associated with an arbitrary secondary reinforcer such as vocal praise; and variable or partial conditioning, where once the correct response has been learnt, reinforcement is presented according to an intermittent schedule to increase resistance to extinction outside of training.

Animals↗

A primacy effect in monkeys when list position is relevant.

In Experiment 1 (1a and 1b), Rhesus monkeys (Macaca mulatta) learned lists of two-choice visual discriminations in which list position was relevant to discrimination performance. For example, Stimulus A was the rewarded stimulus if it was presented at List Position 1, but was not rewarded if it was presented at any other position in the list; similarly, Stimulus B was rewarded only at List Position 2, and so on. In learning these lists, all animals showed a marked primacy effect. In Experiment 2 (2a and 2b), Rhesus monkeys and Cynomolgus monkeys (M. fascicularis) learned lists of visual discriminations in which each visual stimulus occupied a fixed position in a list, but list position was not relevant to discrimination performance. For example, Stimulus E was always rewarded, and was always presented at List Position 1. To increase the salience of list beginning as a distinctive event, successive presentations of the list were separated by 24-hr intervals. In Experiment 2 there was no primacy effect, however. These results show for the first time that a primacy effect can be obtained in visual discrimination learning by monkeys. Furthermore, they suggest that it is obtained only when list position is relevant to the discrimination learning task.

Animals↗

Learning new sounds of speech: reallocation of neural substrates.

Functional magnetic resonance imaging (fMRI) was used to investigate changes in brain activity related to phonetic learning. Ten monolingual English-speaking subjects were scanned while performing an identification task both before and after five sessions of training with a Hindi dental-retroflex nonnative contrast. Behaviorally, training resulted in an improvement in the ability to identify the nonnative contrast. Imaging results suggest that the successful learning of a nonnative phonetic contrast results in the recruitment of the same areas that are involved during the processing of native contrasts, including the left superior temporal gyrus, insula-frontal operculum, and inferior frontal gyrus. Additionally, results of correlational analyses between behavioral improvement and the blood-oxygenation-level-dependent (BOLD) signal obtained during the posttraining Hindi task suggest that the degree of success in learning is accompanied by more efficient neural processing in classical frontal speech regions, and by a reduction of deactivation relative to a noise baseline condition in left parietotemporal speech regions.

Adult↗

Ingestional aversion learning in preweanling rats.

Ingestional aversions were conditioned in 12- and 15-day-old rats by infusing a .5% solution of saccharin into the oral cavity and following this oral infusion by the injection of lithium chloride. At both ages, subjects for which the saccharin exposure was followed by lithium injection within 2-3 min drank less when the saccharin solution was again presented by oral infusion 12 hr later; such suppressions of intake were not observed in subjects that previously received the saccharin and lithium in an unpaired fashion (Experiments 1 and 3). Ingestional aversions were also learned by 12-day-olds when a 30-min interval was introduced between saccharin exposure and lithium toxicosis but not when toxicosis was delayed by 120 min (Experiment 2). In contrast, 15-day-olds learned aversions with both the 30- and 120-min-delay intervals (Experiment 3). Despite the absence of long-delay learning in 12 day olds, ingestional aversions conditioned at 12 days of age were retained for 2 wk (Experiment 4). These results provide further evidence of the associative abilities of neonatal rats and illustrate a developmental aspect of long-delay learning.

Age Factors↗

Different attentional resources modulate the gain mechanisms for color and luminance contrast.

We used an interference paradigm to investigate whether attention is attribute-specific at early levels of visual processing. We show that the peripheral increment thresholds for luminance contrast deteriorate when the observer is currently performing another luminance (form or contrast) discrimination task in central view, but not when he or she is performing a color discrimination task. Similar results were obtained for color increment thresholds, indicating that the interference is specific to contrast modality. The effects are strong and robust over primary task difficulties and perceptual learning levels. Modeling suggests that attention improves contrast sensitivity by modulating the gain of the neuronal response to contrast. These results suggest that attention is parceled in independent resources for luminance and color contrast.

Attention↗

Decreased odor avoidance after electric shock in Drosophila mutants biases learning and memory tests.

The Drosophila mutants amnesiac, dunce (dnc), and rutabaga were isolated after associative conditioning tests, during which animals were trained to associate the presence of an odor with that of electric shocks (ES). In the absence of conditioning, the odor avoidance (OA) of these mutants was shown to be normal, indicating that their poor associative conditioning performance was attributable to specific learning or memory deficits. However, I show that the OA of the mutants is greatly decreased after their exposure to ES. This effect can last for hours. These results strongly suggest that part of the defect displayed by these mutants in associative conditioning tests does not correspond to a learning or memory deficit but might arise from abnormal sensitivity to stressful stimuli. I looked at the OA after ES of two previously characterized dnc mutants. Df(1)N79f specifically decreases Dnc expression in the mushroom bodies, leading to a normal level of learning but decreased memory. Df(1)N79f mutants displayed a normal OA after ES. Df(1)N64j15 affects the entire brain expression of Dnc, leading to decreased learning and memory. Df(1)N64j15 animals showed a strong decrease of their OA after ES. Thus, the lack of Dnc "general" expression is most likely responsible for the OA defect, which would be responsible for the apparent learning defect after conditioning. In contrast, the Dnc phosphodiesterase accumulated in the mushroom bodies would be involved specifically in memory formation.

Amnesia↗

Understanding spatial relations: flexible infants, lexical adults.

Concepts of containment, support, and degree of fit were investigated using nonverbal, preferential-looking tasks with 9- to 14-month-old infants and adults who were fluent in either English or Korean. Two contrasts were tested: tight containment vs. loose support (grammaticized as 'in' and 'on' in English by spatial prepositions and 'kkita' and 'nohta' in Korean by spatial verbs) and tight containment vs. loose containment (both grammaticized as 'in' in English but separately as 'kkita' and 'nehta' in Korean). Infants categorized both contrasts, suggesting conceptual readiness for learning such spatial semantics in either language. English-speaking adults categorized tight containment vs. loose support, but not tight vs. loose containment. However, Korean-speaking adults were successful at this latter contrast, which is lexicalized in their language. The adult data suggest that some spatial relations that are salient during the preverbal stage become less salient if language does not systematically encode them.

Adult↗

Further tests of the Scalar Expectancy Theory (SET) and the Learning-to-Time (LeT) model in a temporal bisection task.

To contrast two models of timing, Scalar Expectancy Theory (SET) and Learning to Time (LeT), pigeons were exposed to a double temporal bisection procedure. On half of the trials, they learned to choose a red key after a 1s signal and a green key after a 4s signal; on the other half of the trials, they learned to choose a blue key after a 4-s signal and a yellow key after a 16-s signal. This was Phase A of an ABA design. On Phase B, the pigeons were divided into two groups and exposed to a new bisection task in which the signals ranged from 1 to 16s and the choice keys were blue and green. One group was reinforced for choosing blue after 1-s signals and green after 16-s signals and the other group was reinforced for the opposite mapping (green after 1-s signals and blue after 16-s signals). Whereas SET predicted no differences between the groups, LeT predicted that the former group would learn the new discrimination faster than the latter group. The results were consistent with LeT. Finally, the pigeons returned to Phase A. Only LeT made specific predictions regarding the reacquisition of the four temporal discriminations. These predictions were only partly consistent with the results.

Algorithms↗

Generalization of learning in three-and-a-half-month-old infants on the basis of amodal relations.

Infants of 3.5 months (N = 124) were given the opportunity to learn to relate two objects and their natural, distinctive sounds during a training phase. The objects and sounds were united by temporal synchrony and amodal audiovisual information specifying object composition. Infants then participated in one of three types of transfer tests (requiring low, moderate, or high degrees of generalization) to measure the extent to which intermodal knowledge generalized to a new task and across events (familiar events; change in color/shape; change in substance, motion, and color/shape). Results indicated that infants tested with the familiar events and with events of a new color/shape showed learning and transfer of knowledge. In contrast, infants tested with events of a new substance, motion, and color/shape showed no generalization of learning. Thus, infants of 3.5 months appear to show a moderate degree of generalization of intermodal knowledge across events. Although this knowledge is not restricted to the events of original learning, it cannot yet be flexibly extended across a variety of contexts.

Auditory Perception↗

Critical flicker frequency series effect in epileptic and in neurotic patients. II. Difference in the variation range ratios of continuous and intermittent CFF series.

In a previous study (1970) a lowering of the CFFSE (Critical Flicker Frequency Series Effect) level from morning to noon was shown. In a group of 18 neurotic subjects 17--34 years of age, an opposite learning effect in the CFFSE level variation, increasing from the first day of examinations to the third one and in a group of 17 epileptic subjects of similar age a decrease of the same was observed. The recent study has corroborated the findings in 195 neurotic and 73 epileptic patients of the same age groups with the following additions. The "negative" learning effect of the epileptics proved to be more intensive in females than in males. The contrast in both positive and negative learning effects between neurotic and epileptic males of 17--34 age was most accentuated in the forenoon. Deviations in the variation range ratios of continuous and intermittent CFF series between the two groups occurred in each age group. The average variation of this ratio in 855 subjects with neurosis, epilepsy and some other diseases seems to point to the occurrence of systematic differences in the pathophysiological function of the brain, changing from organic cerebral lesions through neurosis to organ neuroses.

Adolescent↗

Nucleus basalis magnocellularis lesions facilitate two-way active avoidance.

Bilateral ibotenic acid lesions of the nucleus basalis magnocellularis (NBM), which resulted in 30% depletion of cortical choline acetyltransferase, facilitated acquisition of shuttlebox two-way active avoidance learning in rats. These results contrast with a report of impaired two-way active avoidance after bilateral electrolytic lesions of the NBM, but support findings of facilitation after ibotenic acid lesions. The data suggest that electrolytic lesions impair shuttlebox learning by damaging fibers of passage in the vicinity of the NBM.

Acetylcholinesterase↗

Effects of scopolamine on delayed-matching-to-sample and paired associates tests of visual memory and learning in human subjects: comparison with diazepam and implications for dementia.

Two experiments examined dose-related effects of 200, 400 and 600 micrograms scopolamine (n = 24, s.c.) and 5 and 10 mg diazepam (n = 6, PO) on parallel tests of visual memory and learning taken from the CANTAB battery. Scopolamine significantly impaired accuracy of performance on a delayed matching to sample test of visual recognition memory in a dose- and delay-dependent manner, but had only marginal decremental effects on a test of visuospatial paired associates learning. Scopolamine significantly lengthened decision times in a visual search matching to sample task at the 400 and 600 micrograms doses, without significantly affecting accuracy. The drug also impaired performance on tests of spatial (on accuracy and response time measures) and pattern (on response time only) memory. Most of the deleterious effects on scopolamine were removed by covariance analyses with indices of subjective sedation, but the effects of delayed matching accuracy and latency remained. By contrast, diazepam significantly impaired paired associates learning but affected delayed matching to sample in a delay-independent manner. These results suggest that scopolamine can produce selective deficits in tests of short-term visual recognition memory which do not depend on overall impairments in arousal and which contrast with deficits in visual associative learning produced by diazepam. They have implications for the pharmacological modelling of dementia and memory disorders in man and for the neurochemical substrates of the short-term recognition memory and associative learning for visual stimuli.

Adult↗

Learned helplessness: validity and reliability of depressive-like states in mice.

The learned helplessness paradigm is a depression model in which animals are exposed to unpredictable and uncontrollable stress, e.g. electroshocks, and subsequently develop coping deficits for aversive but escapable situations (J.B. Overmier, M.E. Seligman, Effects of inescapable shock upon subsequent escape and avoidance responding, J. Comp. Physiol. Psychol. 63 (1967) 28-33 ). It represents a model with good similarity to the symptoms of depression, construct, and predictive validity in rats. Despite an increased need to investigate emotional, in particular depression-like behaviors in transgenic mice, so far only a few studies have been published using the learned helplessness paradigm. One reason may be the fact that-in contrast to rats (B. Vollmayr, F.A. Henn, Learned helplessness in the rat: improvements in validity and reliability, Brain Res. Brain Res. Protoc. 8 (2001) 1-7)--there is no generally accepted learned helplessness protocol available for mice. This prompted us to develop a reliable helplessness procedure in C57BL/6N mice, to exclude possible artifacts, and to establish a protocol, which yields a consistent fraction of helpless mice following the shock exposure. Furthermore, we validated this protocol pharmacologically using the tricyclic antidepressant imipramine. Here, we present a mouse model with good face and predictive validity that can be used for transgenic, behavioral, and pharmacological studies.

Animals↗

Context-independent directional cue learning by hippocampal place cells.

In a symmetrical environment possessing no other polarizing visual cues, the spatially localized firing of hippocampal place cells can be primarily orientated by a reliable distal visual stimulus, such as a white cue card. However, if such a directional cue is made unreliable by being frequently moved in full view of the rat, the rat's internal sense of direction comes, over the course of a few days, to control the orientation of place fields instead. We investigated whether this simple form of 'cue-instability' learning would transfer to a new context, in which the firing patterns of the place cells become reorganized and in which a new spatial representation is thus active. We found that after cue-instability learning, the 'remapped' place field representation in the new environment was also orientated by the internal sense of direction of the rat rather than by the cue card, showing that the cue learning generalized from one context (and hence spatial representation) to another. This contrasts with another kind of place cell learning, in which the cells can acquire the ability to discriminate two spatial locations in one context but do not transfer this discrimination to a new context. We discuss the different effects of context changes on learned place cell activity in terms of the possible architecture of the inputs to place cells.

Animals↗

Motivational effects of alcohol on memory consolidation and heart rate in social drinkers.

Several studies have documented the retrograde facilitation of memory by alcohol, but the mechanisms responsible for this curious effect are unknown. In an experiment designed to complement previous studies on incidental learning, social drinkers (men aged 18 to 30; n = 44) took part in an experiment examining the effects of alcohol on intentional learning of emotionally salient verbal stimuli. Learning occurred when participants were sober. Alcohol or placebo (1.0 vs. 0.1 ml/kg) was consumed after learning, and memory was tested, sober, 24 hr later. Compared with placebo, alcohol modestly enhanced recall of positive but not negative stimuli. Furthermore, results suggest that the reinforcing effects on memory for positive (relative to negative) stimuli occurred in association with acute psychomotor stimulant effects during the ascending limb of the blood alcohol curve. The present finding that alcohol appeared to enhance intentional learning in association with its incentive effects contrasts results from previous studies, demonstrating that alcohol appears to enhance incidental learning by memory mechanisms independent of its incentive effects. These findings support a theory of alcoholism that is based on motivational systems.

Adolescent↗

Type of prenatal sensory experience affects prenatal auditory learning in bobwhite quail (Colinus virginianus).

This study examined the effects of specific types of prenatal auditory stimulation on the auditory learning capacity of bobwhite embryos (Colinus virginianus) incubated in either communal or isolation conditions. Results revealed that socially incubated embryos could learn an individual bobwhite maternal call, whereas embryos denied physical and tactile stimulation as a result of isolation incubation failed to demonstrate prenatal auditory learning of the maternal call. In contrast, embryos exposed to bobwhite chick contentment calls in the period prior to hatching demonstrated prenatal auditory learning, whether they were incubated socially or in isolation. Socially incubated and isolation-incubated embryos exposed to bobwhite chick distress calls failed to learn the individual maternal call, indicating that the type of sensory stimulation the developing organism encounters prenatally is important in fostering normal perceptual learning ability.

Animals↗

[Possible mechanisms in latent learning formation investigated by using mutant mice].

We examined possible mechanisms in the development of latent learning by methods of behavioral pharmacology and confirmed them by using mutant mice. Mice that received dopamine agonists, a noradrenergic neurotoxin or a traumatic brain injury showed impairment of latent learning. This impairment was suggested to be mediated by imbalance of dopaminergic and noradrenergic systems since the impairment was attenuated by a noradrenaline uptake inhibitor or a dopamine-D2 antagonist. The heterozygous mice for the tyrosine hydroxylase (TH) gene and for the cyclic AMP (cAMP) response element binding protein (CREB) binding protein (CBP) gene showed impairment of latent learning in the water finding task. The spatial learning and hippocampal long-term potentiation (LTP) were normal in both the mutants. TH heterozygous mice showed a reduction of high K(+)-evoked noradrenaline release in the frontal cortex by the microdialysis technique and a reduction of cAMP of the brain cAMP content. The central noradrenergic systems and/or cAMP signal pathways play an important role in latent learning, but not spatial memory. In contrast with TH and CBP mutant mice, nociceptin-knockout mice showed an enhanced retention of latent learning in the water finding task, greater learning ability in the water maze task and larger LTP than wild-type mice. Such mice showed hyperfunction of dopaminergic systems in the cortex. Nociceptin itself induced latent learning impairment in wild-type mice. These results suggest that the nociceptin system seems to play negative roles in learning and memory. In conclusion, the results of mutant mice further supported our previous results of behavioral pharmacology and suggest that the alternation of catecholamine biosynthesis and cAMP signal pathways may play a key role in development of latent learning. They further suggest that the expression of genes mediated by phosphorylated CREB may be involved in the development of latent learning.

Adenylyl Cyclases↗