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Within-compound associations in retrospective revaluation and in direct learning: a challenge for comparator theory.

In three human causal learning experiments we investigated the role of within-compound associations in learning about absent cues versus learning about present cues. Different theoretical approaches agree that within-compound associations are essential for learning about absent cues-that is, for retrospective revaluation. They differ, however, with regard to the role of within-compound associations for learning about present cues-that is, for direct learning. A memory test was used to assess within-compound associations. Experiment 1 used a blocking/release from overshadowing design, Experiment 2 used a conditioned inhibition design, and Experiment 3 used a higher-order cue selection design. In all experiments, first-order retrospective revaluation was significantly correlated with within-compound associations, but no significant correlations were found for the direct learning conditions. In addition to this, second-order retrospective revaluation in Experiment 3 was positively correlated to joint knowledge of first-order and second-order within-compound associations. Furthermore, cue selection effects were stronger for direct learning conditions than for retrospective learning conditions. These results are at variance with the comparator hypothesis but are in agreement with a modified associative theory and with the suggestion that retrospective revaluation might be due to rehearsal processes.

Adolescent↗

Aging and probabilistic learning in single- and multiple-cue tasks.

The objective of the study was to examine some of the conditions under which elderly people are able to learn probabilistic inverse relationships and when this type of learning is no longer possible. Two kind of tasks were used: (a) two single-cue learning tasks with either direct or inverse relationships (the Single-Cue Probability Learning paradigm), and (b) three two-cue learning tasks, one with two direct relationships, one with a combination of direct and inverse relationships, and one with two inverse relationships (the Multiple-Cue Probability Learning paradigm). Four groups of participants were included in the study: young adults (18- to 25-year-olds), adults (40- to 50-year-olds), elderly people (65- to 74-year-olds), and very elderly people (75- to 90-year-olds). It was shown that (a) older adults are able to reject the direct relationship "default" hypothesis and select the inverse relationship hypothesis when outcome feedback contradicting the default hypothesis is given, and provided that the learning setting be a very simple one, involving only one cue; (b) some older adults are able to select the inverse relationship hypothesis provided that the learning setting be a simple one,involving only two inverse relationship cues; and (c) very few older adults are able to select the inverse relationship hypothesis when the learning setting is a complex one, involving two cues with both direct and inverse relationships with the criterion. These results led to the revision of the "gradual decrease of cognitive flexibility in older adults" hypothesis proposed by Chasseigne, Mullet, and Stewart (Acta Psychologicgrave;a, 103, 229-238, 1997).

Adolescent↗

The implicit benefit of learning without errors.

Two studies examined whether the number of errors made in learning a motor skill, golf putting, differentially influences the adoption of a selective (explicit) or unselective (implicit) learning mode. Errorful learners were expected to adopt an explicit, hypothesis-testing strategy to correct errors during learning, thereby accruing a pool of verbalizable rules and exhibiting performance breakdown under dual-task conditions, characteristic of a selective mode of learning. Reducing errors during learning was predicted to minimize the involvement of explicit hypothesis testing leading to the adoption of an unselective mode of learning, distinguished by few verbalizable rules and robust performance under secondary task loading. Both studies supported these predictions. The golf putting performance of errorless learners in both studies was unaffected by the imposition of a secondary task load, whereas the performance of errorful learners deteriorated. Reducing errors during learning limited the number of error-correcting hypotheses tested by the learner, thereby reducing the contribution of explicit processing to skill acquisition. It was concluded that the reduction of errors during learning encourages the use of implicit, unselective learning processes, which confer insusceptibility to performance breakdown under distraction.

Adult↗

Mnemonic strategies in older people: a comparison of errorless and errorful learning.

OBJECTIVE: To compare the efficacy of errorless and errorful learning on memory performance in older people and young adults. METHODS: Face-name association learning was examined in 18 older people and 16 young controls. Subjects were either prompted to guess the correct name during the presentation of photographs of unknown faces (errorful learning) or were instructed to study the face without guessing (errorless learning). The correct name was given after the presentation of each face in both task conditions. Uncued testing followed immediately after the two study phases and after a 10-minute delay. RESULTS: Older subjects had an overall lower memory performance and flatter learning curves compared to the young adults, regardless of task conditions. Also, errorless learning resulted in a higher accuracy than errorful learning, to an equal amount in both groups. CONCLUSIONS: Older people have difficulty in the encoding stages of face-name association learning, whereas retrieval is relatively unaffected. In addition, the prevention of errors occurring during learning results in a better memory performance, and is perhaps an effective strategy for coping with age-related memory decrement.

Adult↗

PET activation of the medial temporal lobe in learning.

Regional cerebral blood flow was examined during multiple-trial learning in healthy volunteers. On the basis that incremental learning from trial to trial is severely impaired in neuropsychological studies of patients with medial temporal lesions, we predicted that medial temporal activation might be particularly associated with incremental gains in learning. On the other hand, we predicted that frontal activations would not show any increase during incremental learning, and might even diminish. PET recordings were undertaken while subjects were presented visually with a 15-word list in one of three conditions: a list in which a single word was repeated 15 times (S), a list of novel words (N), and a list which was repeated from before (R). We demonstrated that statistically significant incremental learning did occur when word lists were repeated in (R) trials. The subtraction of novel minus repeated conditions (N-R) was associated with left medial temporal as well as left prefrontal activations, whereas the opposite (R-N) subtraction gave rise to right prefrontal and precuneal activations. In particular, incremental learning during the repeated trials (R) identified a left medial temporal activation, as predicted, but the left frontal activation was no longer evident. We suggest that the left medial temporal region is not only activated by novel, to-be-learned stimuli, but it also contributes to incremental learning as part of a network involved in 'binding' or 'consolidating' new memories. The right frontal and precuneal regions, which participate in the repeated retrieval and rehearsal of already learned memories, are also involved in this network. The left frontal region is implicated in the more 'effortful' or elaborative aspects of memory.

Adult↗

How medical students learn.

With recent concerns over and attempts to address issues regarding medical students' processes of learning and development of lifelong learning habits, a question often asked is: How do medical students learn in the present medical curriculum? In this study, the authors directly attempt to answer that question--they assess the learning behaviors of 254 students in two medical schools, one with an objectives-based mastery and one with a traditional curricula, and discuss whether the students' adopted behaviors are conducive to the development of effective, analytic, and independent learning. Most students of both medical schools reported very similar effective and ineffective learning behaviors. The students' behaviors were found to dispose them more to analytic learning than to independent learning. Suggestions to restructure the curriculum for better development of students' independent learning habits are proposed.

Behavior↗

Rapid word-learning in normal-hearing and hearing-impaired children: effects of age, receptive vocabulary, and high-frequency amplification.

OBJECTIVE: This study examined rapid word-learning in 5- to 14-year-old children with normal and impaired hearing. The effects of age and receptive vocabulary were examined as well as those of high-frequency amplification. Novel words were low-pass filtered at 4 kHz (typical of current amplification devices) and at 9 kHz. It was hypothesized that (1) the children with normal hearing would learn more words than the children with hearing loss, (2) word-learning would increase with age and receptive vocabulary for both groups, and (3) both groups would benefit from a broader frequency bandwidth. DESIGN: Sixty children with normal hearing and 37 children with moderate sensorineural hearing losses participated in this study. Each child viewed a 4-minute animated slideshow containing 8 nonsense words created using the 24 English consonant phonemes (3 consonants per word). Each word was repeated 3 times. Half of the 8 words were low-pass filtered at 4 kHz and half were filtered at 9 kHz. After viewing the story twice, each child was asked to identify the words from among pictures in the slide show. Before testing, a measure of current receptive vocabulary was obtained using the Peabody Picture Vocabulary Test (PPVT-III). RESULTS: The PPVT-III scores of the hearing-impaired children were consistently poorer than those of the normal-hearing children across the age range tested. A similar pattern of results was observed for word-learning in that the performance of the hearing-impaired children was significantly poorer than that of the normal-hearing children. Further analysis of the PPVT and word-learning scores suggested that although word-learning was reduced in the hearing-impaired children, their performance was consistent with their receptive vocabularies. Additionally, no correlation was found between overall performance and the age of identification, age of amplification, or years of amplification in the children with hearing loss. Results also revealed a small increase in performance for both groups in the extended bandwidth condition but the difference was not significant at the traditional p = 0.05 level. CONCLUSIONS: The ability to learn words rapidly appears to be poorer in children with hearing loss over a wide range of ages. These results coincide with the consistently poorer receptive vocabularies for these children. Neither the word-learning or receptive-vocabulary measures were related to the amplification histories of these children. Finally, providing an extended high-frequency bandwidth did not significantly improve rapid word-learning for either group with these stimuli.

Adolescent↗

Towards an explicit account of implicit learning.

PURPOSE OF REVIEW: The human brain supports acquisition mechanisms that can extract structural regularities implicitly from experience without the induction of an explicit model. Reber defined the process by which an individual comes to respond appropriately to the statistical structure of the input ensemble as implicit learning. He argued that the capacity to generalize to new input is based on the acquisition of abstract representations that reflect underlying structural regularities in the acquisition input. We focus this review of the implicit learning literature on studies published during 2004 and 2005. We will not review studies of repetition priming ('implicit memory'). Instead we focus on two commonly used experimental paradigms: the serial reaction time task and artificial grammar learning. Previous comprehensive reviews can be found in Seger's 1994 article and the Handbook of Implicit Learning. RECENT FINDINGS: Emerging themes include the interaction between implicit and explicit processes, the role of the medial temporal lobe, developmental aspects of implicit learning, age-dependence, the role of sleep and consolidation. SUMMARY: The attempts to characterize the interaction between implicit and explicit learning are promising although not well understood. The same can be said about the role of sleep and consolidation. Despite the fact that lesion studies have relatively consistently suggested that the medial temporal lobe memory system is not necessary for implicit learning, a number of functional magnetic resonance studies have reported medial temporal lobe activation in implicit learning. This issue merits further research. Finally, the clinical relevance of implicit learning remains to be determined.

Brain↗

Motor learning: its relevance to stroke recovery and neurorehabilitation.

PURPOSE OF REVIEW: Much of neurorehabilitation rests on the assumption that patients can improve with practice. This review will focus on arm movements and address the following questions: (i) What is motor learning? (ii) Do patients with hemiparesis have a learning deficit? (iii) Is recovery after injury a form of motor learning? (iv) Are approaches based on motor learning principles useful for rehabilitation? RECENT FINDINGS: Motor learning can be broken into kinematic and dynamic components. Studies in healthy subjects suggest that retention of motor learning is best accomplished with variable training schedules. Animal models and functional imaging in humans show that the mature brain can undergo plastic changes during both learning and recovery. Quantitative motor control approaches allow differentiation between compensation and true recovery, although both improve with practice. Several promising new rehabilitation approaches are based on theories of motor learning. These include impairment oriented-training (IOT), constraint-induced movement therapy (CIMT), electromyogram (EMG)-triggered neuromuscular stimulation, robotic interactive therapy and virtual reality (VR). SUMMARY: Motor learning mechanisms are operative during spontaneous stroke recovery and interact with rehabilitative training. For optimal results, rehabilitation techniques should be geared towards patients' specific motor deficits and possibly combined, for example, CIMT with VR. Two critical questions that should always be asked of a rehabilitation technique are whether gains persist for a significant period after training and whether they generalize to untrained tasks.

Electric Stimulation Therapy↗

Smells are no surer: rapid improvement in olfactory discrimination is not due to the acquisition of a learning set.

The claim that rats can demonstrate the 'primate-like' learning capacity of learning set formation when trained with olfactory cues, rather than visual or auditory cues, has generated considerable interest in recent years. In this study, the claim is evaluated in detail by using a series of experimental and control procedures to determine whether rats do indeed develop the abstract 'win-stay, lose-shift' strategy which underlies learning set formation in monkeys. We report here that although exposure to a series of novel olfactory discrimination problems gives rise to progressive improvement in the rate of learning, it is not a necessary condition for the development of that rapid learning. Furthermore, even rats which fail to display progressive improvement in olfactory reversal learning show rapid learning on novel olfactory discrimination problems. Each of these findings suggests that although olfactory learning may be rapid, learning set formation does not occur over a short series of problems.

Animals↗

Quality of song learning affects female response to male bird song.

Bird song is unusual as a sexually selected trait because its expression depends on learning as well as genetic and other environmental factors. Prior work has demonstrated that males who are deprived of the opportunity to learn produce songs that function little if at all in male-female interactions. We asked whether more subtle variation in male song-learning abilities influences female response to song. Using a copulation solicitation assay, we measured the response of female song sparrows (Melospiza melodia) to songs of laboratory-reared males that differed in the amount of learned versus invented material that they included and in the degree to which learned material accurately matched the model from which it was copied. Females responded significantly more to songs that had been learned better, by either measure. Females did not discriminate between the best-learned songs of laboratory-reared males and songs of wild males used as models during learning. These results provide, to our knowledge, a first experimental demonstration that variation in learning abilities among males plays a functionally important part in the expression of a sexually selected trait, and further provide support for the hypothesis that song functions as an indicator of male quality because it reflects variation in response to early developmental stress.

Animals↗

Hippocampal synaptic plasticity: role in spatial learning or the automatic recording of attended experience?

Allocentric spatial learning can sometimes occur in one trial. The incorporation of information into a spatial representation may, therefore, obey a one-trial correlational learning rule rather than a multi-trial error-correcting rule. It has been suggested that physiological implementation of such a rule could be mediated by N-methyl-D-aspartate (NMDA) receptor-dependent long-term potentiation (LTP) in the hippocampus, as its induction obeys a correlational type of synaptic learning rule. Support for this idea came originally from the finding that intracerebral infusion of the NMDA antagonist AP5 impairs spatial learning, but studies summarized in the first part of this paper have called it into question. First, rats previously given experience of spatial learning in a watermaze can learn a new spatial reference memory task at a normal rate despite an appreciable NMDA receptor blockade. Second, the classical phenomenon of 'blocking' occurs in spatial learning. The latter finding implies that spatial learning can also be sensitive to an animal's expectations about reward and so depend on more than the detection of simple spatial correlations. In this paper a new hypothesis is proposed about the function of hippocampal LTP. This hypothesis retains the idea that LTP subserves rapid one-trial memory, but abandons the notion that it serves any specific role in the geometric aspects of spatial learning. It is suggested that LTP participates in the automatic recording of attended experience': a subsystem of episodic memory in which events are temporarily remembered in association with the contexts in which they occur. An automatic correlational form of synaptic plasticity is ideally suited to the online registration of context event associations. In support, it is reported that the ability of rats to remember the most recent place they have visited in a familiar environment is exquisitely sensitive to AP5 in a delay-dependent manner. Moreover, new studies of the lasting persistence of NMDA-dependent LTP, known to require protein synthesis, point to intracellular mechanisms that enable transient synaptic changes to be stabilized if they occur in close temporal proximity to important events. This new property of hippocampal LTP is a desirable characteristic of an event memory system.

2-Amino-5-phosphonovalerate↗

Learning "what" and "how" in a human motor task.

We studied the development of implicit and of verbally declared knowledge for normal human subjects who learned an unfamiliar motor task in one learning session. The exploratory nature of motor learning and a special period for optimizing skill were followed in real time. Subjects understood the goal for task success, but they had to learn a motor strategy of what pattern of serial movements to make and the tactics of how much to scale their amplitudes and timing. We compared the time course for acquiring tactical skill with that for acquiring knowledge of strategy and of tactics, and their necessary cues. Implicit and declarative knowledge were distinguished from one another by correlating subjects' verbal self-reports with movement kinematics and their results. Implicit generation of the correct strategy and of the tactics developed in an exploratory manner from the beginning of the learning session. Implicit strategy learning soon gave way to conscious efforts, but tactical learning remained implicit until its first unambiguous verbal declaration (with one exception). First strategy declarations were voiced before those for tactics, during trial-and-error learning that did not require task success, and referred to reversing the direction of hand movements (one exception). In contrast, first declarations of tactics almost always required actual or imminent success, referred to when direction was to be reversed, and it was achieved near the top of a sigmoid learning curve that rose to tactical skill (with one exception). During the sigmoid rise, movement amplitudes and timing were optimized in a distinct manner, although tactics usually adapted thereafter to movements of more moderate speed that could still be successful.

Adult↗

Polysynaptic potentiation at different levels of rat olfactory pathways following learning.

This study was aimed at investigating the consequences of learning on late polysynaptic components of evoked field potential signals recorded in parallel at different levels of the olfactory pathways. For this, evoked field potentials induced by electrical stimulation of the olfactory bulb were recorded simultaneously in the anterior piriform cortex, the posterior piriform cortex, the lateral entorhinal cortex, and the dentate gyrus. The different parameters of late components were measured in each site before and after completion of associative learning in anesthetized rats. In the learning task, rats were trained to associate electrical stimulation of one olfactory bulb electrode with the delivery of sucrose (positive reward) and stimulation of a second olfactory bulb electrode with the delivery of quinine (negative reward). In this way, stimulation of the same olfactory bulb electrodes used for inducing field potentials served as a discriminative cue in the learning paradigm. The data confirmed previous observation that learning was associated with a lowering in late-component-1 intensity of induction in the posterior piriform cortex. The use of simultaneous recording allowed us to further specify the consequences of learning on late-component distribution in the studied network. Indeed the data showed that whereas before learning, late component 1 was rather uniformly distributed among the recorded sites; following learning, its expression was facilitated preferentially in the posterior piriform cortex and lateral entorhinal cortex. Furthermore, learning was accompanied by the emergence of a new late component (late component 2), which occurred simultaneously in the four recording sites. The possible involvement of potentiation of polysynaptic components in recognition and/or consolidation processes will be discussed.

Animals↗

Sleep and the time course of motor skill learning.

Growing evidence suggests that sleep plays an important role in the process of procedural learning. Most recently, sleep has been implicated in the continued development of motor-skill learning following initial acquisition. However, the temporal evolution of motor learning before and after sleep, the effects of different training regimens, and the long-term development of motor learning across multiple nights of sleep remain unknown. Here, we report data for subjects trained and retested on a sequential finger-tapping task across multiple days. The findings demonstrate firstly that following initial training, small practice-dependent improvements are possible before, but not following the large practice-independent gains that develop across a night of sleep. Secondly, doubling the quantity of initial training does not alter the amount of subsequent sleep-dependent learning that develops overnight. Thirdly, the amount of sleep-dependent learning does not correlate with the amount of practice-dependent learning achieved during training, suggesting the existence of two discrete motor-learning processes. Finally, whereas the majority of sleep-dependent motor-skill learning develops during the first night of sleep following training, additional nights of sleep still offer continued improvements.

Adult↗

Intrahippocampal muscimol shifts learning strategy in gonadally intact young adult female rats.

Learning strategy preferences depend upon circulating estrogen levels, with enhanced hippocampus-sensitive place learning coinciding with elevated estrogen levels. The effects of estrogen on strategy may be mediated by fluctuations in GABAergic function, given that inhibitory tone in the hippocampus is low when estrogen is high. We investigated the effects on learning strategy of intrahippocampal injections of a GABA(A) agonist in gonadally intact female rats. On the day of training, rats received 0.3 microL intrahippocampal infusions of muscimol (0.26 nmol or 2.6 nmol) or saline 20 min prior to training on a T-maze in which place (hippocampus-sensitive) or response (striatum-sensitive) strategies offer effective solutions. Muscimol treatment increased the use of the response strategy in a dose-dependent manner without influencing learning speed, indicating that muscimol modulated strategy and not learning ability. Furthermore, the muscimol-related shift to response strategies varied across the estrous cycle. The results indicate that increasing inhibition in the hippocampus biases rats away from hippocampus-sensitive place learning strategies and toward hippocampus-insensitive response learning strategies without a learning deficit. Furthermore, rats at proestrus demonstrated the most dramatic shift in learning strategy following muscimol treatment compared with control conditions, while rats at estrus demonstrated the most complete bias toward response strategies. The enhanced use of hippocampus-sensitive strategies at proestrus likely results from reduced hippocampal inhibition.

Animals↗

Song learning accelerates allopatric speciation.

The songs of many birds are unusual in that they serve a role in identifying conspecific mates, yet they are also culturally transmitted. Noting the apparently high rate of diversity in one avian taxon, the songbirds, in which song learning appears ubiquitous, it has often been speculated that cultural transmission may increase the rate of speciation. Here we examine the possibility that song learning affects the rate of allopatric speciation. We construct a population-genetic model of allopatric divergence that explores the evolution of genes that underlie learning preferences (predispositions to learn some songs over others). We compare this with a model in which mating signals are inherited only genetically. Models are constructed for the cases where songs and preferences are affected by the same or different loci, and we analyze them using analytical local stability analysis combined with simulations of drift and directional sexual selection. Under nearly all conditions examined, song divergence occurs more readily in the learning model than in the nonlearning model. This is a result of reduced frequency-dependent selection in the learning models. Cultural evolution causes males with unusual genotypes to tend to learn from the majority of males around them, and thus develop songs compatible with the majority of the females in the population. Unusual genotypes can therefore be masked by learning. Over a wide range of conditions, learning therefore reduces the waiting time for speciation to occur and can be predicted to accelerate the rate of speciation.

Animals↗

Learning styles and approaches: implications for medical education.

This paper reviews the recent literature on learning styles and approaches to learning. It identifies two separate streams of research, one originating from mainstream cognitive and psychometric psychology and one from research undertaken within the everyday learning environment. The latter is dealt with in greater detail as it seems to have more immediate practical relevance. A simple model of the teaching-learning process is presented showing how students learn in different ways which are partly attributable to their preferred learning style and partly to the context in which the learning takes place. Three basic approaches have been identified: surface, deep and strategic, each resulting in a different learning outcome. The most desirable and successful is the deep approach. The way in which the teaching and the policies of the department and school influence the students' approach to learning are reviewed in some detail. A consideration of these characteristics in medical schools suggested that many may hinder rather than assist in the development of the desired approach. The work reviewed here suggests that the remedy will require not only substantial changes in the teaching, curriculum and, particularly, assessment, but also a new strategy based on identifying and assisting individual students whose approaches to study are not those expected of a competent university-educated doctor.

Australia↗