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Prototype-distortion category learning: a two-phase learning process across a distributed network.

This paper reviews a body of work conducted in our laboratory that applies functional magnetic resonance imaging (fMRI) to better understand the biological response and change that occurs during prototype-distortion learning. We review results from two experiments (Little, Klein, Shobat, McClure, & Thulborn, 2004; Little & Thulborn, 2005) that provide support for increasing neuronal efficiency by way of a two-stage model that includes an initial period of recruitment of tissue across a distributed network that is followed by a period of increasing specialization with decreasing volume across the same network. Across the two studies, participants learned to classify patterns of random-dot distortions (Posner & Keele, 1968) into categories. At four points across this learning process subjects underwent examination by fMRI using a category-matching task. A large-scale network, altered across the protocol, was identified to include the frontal eye fields, both inferior and superior parietal lobules, and visual cortex. As behavioral performance increased, the volume of activation within these regions first increased and later in the protocol decreased. Based on our review of this work we propose that: (i) category learning is reflected as specialization of the same network initially implicated to complete the novel task, and (ii) this network encompasses regions not previously reported to be affected by prototype-distortion learning.

Adult↗

Dissociating explicit and procedural-learning based systems of perceptual category learning.

A fundamental question is whether people have available one category learning system, or many. Most multiple systems advocates postulate one explicit and one implicit system. Although there is much agreement about the nature of the explicit system, there is less agreement about the nature of the implicit system. In this article, we review a dual systems theory of category learning called competition between verbal and implicit systems (COVIS) developed by Ashby et al. The explicit system dominates the learning of verbalizable, rule-based category structures and is mediated by frontal brain areas such as the anterior cingulate, prefrontal cortex (PFC), and head of the caudate nucleus. The implicit system, which uses procedural learning, dominates the learning of non-verbalizable, information-integration category structures, and is mediated by the tail of the caudate nucleus and a dopamine-mediated reward signal. We review nine studies that test six a priori predictions from COVIS, each of which is supported by the data.

Caudate Nucleus↗

Cognitive sequence learning in Parkinson's disease and amnestic mild cognitive impairment: Dissociation between sequential and non-sequential learning of associations.

Evidence suggests that dopaminergic mechanisms in the basal ganglia (BG) are important in the learning of sequential associations. To test the specificity of this hypothesis, we assessed never-medicated patients with Parkinson's disease (PD) and amnestic mild cognitive impairment (aMCI) using a chaining task. In the training phase of the chaining task, each link in a sequence of stimuli leading to reward is trained step-by-step using feedback after each decision, until the complete sequence is learned. In the probe phase of the chaining task, the context of stimulus-response associations must be used (the position of the associations in the sequence). Results revealed that patients with PD showed impaired learning during the training phase of the chaining task, but their performance was spared in the probe phase. In contrast, patients with aMCI with prominent medial temporal lobe (MTL) dysfunctions showed intact learning during the training phase of the chaining task, but their performance was impaired in the probe phase of the chaining task. These results indicate that when dopaminergic mechanisms in the BG are dysfunctional, series of stimulus-response associations are less efficiently acquired, but their sequential manner is maintained. In contrast, MTL dysfunctions may result in a non-sequential learning of associations, which may indicate a loss of contextual information.

Aged↗

Level and mechanisms of perceptual learning: learning first-order luminance and second-order texture objects.

Perceptual learning is an improvement in perceptual task performance reflecting plasticity in the perceptual system. Practice effects were studied in two object orientation tasks: a first order, luminance object task and a second-order, texture object task. Perceptual learning was small or absent in the first-order task, but consistently occurred for the second-order (texture) task, where it was limited to improvements in low external noise conditions, or stimulus enhancement [Dosher, B., & Lu, Z. -L. (1998). Perceptual learning reflects external noise filtering and internal noise reduction through channel reweighting. Proceedings of the National Academy of Sciences of the United States of America, 95 (23) 13988-13993; Dosher, B., & Lu, Z. -L. (1999). Mechanisms of perceptual learning. Vision Research, 39 (19) 3197-3221], analogous to attention effects in first- and second-order motion processing [Lu, Z. -L., Liu, C. Q., & Dosher, B. (2000). Attention mechanisms for multi-location first- and second-order motion perception. Vision Research, 40 (2) 173-186]. Perceptual learning affected the later, post-rectification, stages of perceptual analysis, possibly localized at V2 or above. It serves to amplify the stimulus relative to limiting internal noise for intrinsically noisy representations of second-order stimuli.

Attention↗

Associations between student learning outcomes from their clinical placement and their perceptions of the social climate of the clinical learning environment.

The aim of this study was to investigate the associations between student learning outcomes from their clinical placement and their perceptions of the social climate of the clinical learning environment. The Clinical Learning Environment Inventory (CLEI) was used to collect data. The final sample group consisted of 108 second-year nursing students undertaking clinical placement in 14 metropolitan hospitals in South Australia. Findings from the study suggested that students' perceptions of the outcomes of their clinical placement are strongly associated with all five scales of the CLEI namely; Individualisation, Innovation, Involvement, Personalisation and Task Orientation. It is interesting to note that student who perceived the outcomes of their clinical placement as greatly positive, have placed high expectations on the scale, Task Orientation. The study also found that there were significant differences between students' perceptions of the actual clinical learning environment with their preferred clinical learning environment. The value of this study lies in the resulting implication for nursing education and future research. A better understanding of what constitutes quality clinical education from the students' perspective would be valuable in providing better educational experiences.

Education, Nursing↗

The cerebellum and cognition: cerebellar lesions impair sequence learning but not conditional visuomotor learning in monkeys.

Claims that the cerebellum contributes to cognitive processing in humans have arisen from both functional neuroimaging and patient studies. These claims challenge traditional theories of cerebellar function that ascribe motor functions to this structure. We trained monkeys to perform both a visuomotor conditional associative learning task and a visually guided sequence task, and studied the effects of bilateral excitotoxic lesions in the lateral cerebellar nuclei. In the first experiment three operated monkeys showed a small impairment in post-operative retention of a visuomotor associative task (A) but were then not impaired in learning a new task (B). However, the impairment on A could have been due to a problem in making the movements themselves. In a second experiment we therefore gave the three control animals a further pre-operative retest on both A and B and then tested after surgery on retention of both tasks. Though again the animals showed motor problems on task A, they reached criterion, and at this stage could clearly make both movements satisfactorily. The critical test was then retention of task B, and they were not impaired. In the final experiment (serial reaction time task) the monkeys response times on a repeating visuomotor sequence were compared with those for a pseudo-random control sequence. After bilateral nuclei lesions they were slow to execute the pre-operatively learned sequence but were still faster on this than on the control task. However, when they were then given a new repeating sequence to learn, they never performed the sequence as quickly as they had on retention of the first sequence. We conclude that the cerebellum is not essential for the learning or recall of stimulus-response associations but that it is crucially involved in the process by which motor sequences become automatic with extended practice.

Animals↗

Neurobiology of associative learning in the neonate: early olfactory learning.

Mammalian neonates have been simultaneously described as having particularly poor memory, as evidenced by infantile amnesia, and as being particularly excellent learners with unusually plastic nervous systems that are easily influenced by experience. An understanding of the neurobiological constraints and mechanisms of early learning may contribute to a unified explanation of these two disparate views. Toward that end, we review here our work on the neurobiology of learning and memory in neonates. Specifically, we have examined the neurobiology of early learning using an olfactory classical conditioning paradigm. Olfactory classical conditioning in neonates at the behavioral level conforms well with the requirements and outcomes of classical conditioning described in adults. Furthermore, specific neural correlates of this behavioral conditioning have been described including anatomical and physiological changes, neural pathways, and modulatory systems. In this Review, we outline the behavioral paradigm, the identified neural correlates, and apparent mechanisms of this learning. Finally, we compare the neurobiology of early learning with that reported for mature animals, with specific reference to the role of US-CS convergence, memory modulation, consolidation, and distributed memory.

Animals↗

Enhancement of spatial learning in F344 rats by physical activity and related learning-associated alterations in hippocampal and cortical cholinergic functioning.

The effects of physical activity on spatial memory performance and associated cholinergic function were examined in F344 rats. Cholinergic analysis included resting and depolarization-induced activation of high-affinity choline uptake and muscarinic receptor binding in the hippocampus, parietal cortex and frontal cortex. Rats that were physically trained, using chronic treadmill running, demonstrated significantly enhanced performance on the spatial learning task, both in second trial latency and first and second trial proximity ratio scores (P less than 0.002). Concomitant with enhanced behavioral performance were neurochemical changes of a reduction in hippocampal high-affinity choline uptake, an upregulation of muscarinic receptor density, and an increase in high-affinity choline uptake 24 h after spatial memory testing (P less than 0.05). Spatial memory tested rats demonstrated enhanced depolarization-induced activation of high-affinity choline uptake (P less than 0.001). Rats that were yoked for swim time to spatial memory tested rats did not show any spatial learning-induced alterations in high affinity choline uptake. These spatial learning- and physical activity-induced cholinergic alterations were observed only in the hippocampus, not in the parietal or frontal cortex. These data indicate that the chronic running-induced alterations in hippocampal high-affinity choline uptake and upregulation of muscarinic receptor density, in combination with enhancement of high-affinity choline uptake related to spatial learning, may contribute to the enhanced spatial learning performance of chronic-run rats.

Animals↗

Freedom to enjoy learning in the 21st century: developing an active learning culture in nursing.

In 1994, the Wolfson School of Health Sciences was created in Thames Valley University, UK. This paper discusses an active learning approach being implemented throughout the university, and gives an outline of its application within nursing education. Practising as a confident, competent, reflective practitioner in the profession of nursing cannot automatically commence on the day of registration. Rather, the philosophical underpinnings necessary should be fostered throughout the initial nursing education curricula. In a political climate where there is a strong emphasis on efficiency, nurse teachers need to reconsider how the acquisition of nursing knowledge is facilitated. A description of how an active learning approach is being further developed in the nursing skills laboratory and the physiology laboratory is given. An outline identifies how these facilities may help students of nursing to gain knowledge and skills in relation to respiration. The authors recognize that while laboratories have been used widely in general education, their use has been limited in nursing education, particularly in the UK. Informal feedback from students implies that not only do they enjoy learning in these facilities, but also motivation towards learning is raised, an important element in the learning process. It is suggested that this will result ultimately in increased nursing competence. The authors acknowledge that these are preliminary informal findings and that formal investigation is required.

Education, Nursing, Baccalaureate↗

Brain mechanisms of selective learning: event-related potentials provide evidence for error-driven learning in humans.

Selective learning has been observed in Pavlovian conditioning in animals and in judgements of event contingencies in humans. This analogy led to the suggestion that the formation of associations underlies both types of learning. An alternative theory proposes that both tasks involve the computation of event contingencies as prescribed by probability theory. Error-driven models of learning incorporate trial-by-trial error-correction mechanisms during training whereas probabilistic models view learning merely as the storage of frequency information for later use during judgement of event contingencies. Competitive interaction between cues was observed in a contingency judgement task. Event-related brain potentials (ERPs) provided evidence for brain events related to the discrepancy between actual and expected outcomes during training thus supporting error-driven accounts of selective learning.

Adult↗

Distance learning, problem based learning and dynamic knowledge networks.

This paper is an attempt to develop a distance learning model grounded upon a strict integration of problem based learning (PBL), dynamic knowledge networks (DKN) and web tools, such as hypermedia documents, synchronous and asynchronous communication facilities, etc. The main objective is to develop a theory of distance learning based upon the idea that learning is a highly dynamic cognitive process aimed at connecting different concepts in a network of mutually supporting concepts. Moreover, this process is supposed to be the result of a social interaction that has to be facilitated by the web. The model was tested by creating a virtual classroom of medical and nursing students and activating a learning session on the concept of knowledge representation in health sciences.

Computer Communication Networks↗

Learning to read is much more than learning to read: a neuropsychologically based reading program.

Departing from the observation that illiterates significantly underscore in some neuropsychological tests, a learning-to-read method named NEUROALFA was developed. NEUROALFA is directed to reinforce these underscored abilities during the learning-to-read process. It was administered to a sample of 21 adult illiterates in Colima (Mexico). Results were compared with 2 control groups using more traditional procedures in learning to read. The NEUROPSI neuropsychological test battery was administered to all the participants before and after completing the learning-to-read training program. All 3 groups presented some improvement in the test scores. Gains, however, were significantly higher in the experimental group in Orientation in Time, Digits Backward, Visual Detection, Verbal Memory, Copy of a Semi-Complex Figure, Language Comprehension, Phonological Verbal Fluency, Similarities, Calculation Abilities, Sequences, and all the recall subtests, excluding Recognition. Performance in standard reading tests was also significantly higher in the experimental group. Correlations between pretest NEUROPSI scores and reading ability were low. However, correlations between posttest NEUROPSI scores and reading scores were higher and significant for several subtests. Results are interpreting as supporting the assumption that reinforcement of those abilities in which illiterates significantly underscore results in a significant improvement in neuropsychological test scores and strongly facilitates the learning-to-read process. The NEUROALFA method of teaching reading to adult illiterates is beginning to be used extensively in Mexico. To our knowledge, this is the first attempt to apply neuropsychological principles to social problems.

Adolescent↗

Striatal contributions to category learning: quantitative modeling of simple linear and complex nonlinear rule learning in patients with Parkinson's disease.

The contribution of the striatum to category learning was examined by having patients with Parkinson's disease (PD) and matched controls solve categorization problems in which the optimal rule was linear or nonlinear using the perceptual categorization task. Traditional accuracy-based analyses, as well as quantitative model-based analyses were performed. Unlike accuracy-based analyses, the model-based analyses allow one to quantify and separate the effects of categorization rule learning from variability in the trial-by-trial application of the participant's rule. When the categorization rule was linear, PD patients showed no accuracy, categorization rule learning, or rule application variability deficits. Categorization accuracy for the PD patients was associated with their performance on a test believed to be sensitive to frontal lobe functioning. In contrast, when the categorization rule was nonlinear, the PD patients showed accuracy, categorization rule learning, and rule application variability deficits. Furthermore, categorization accuracy was not associated with performance on the test of frontal lobe functioning. Implications for neuropsychological theories of categorization learning are discussed.

Aged↗

Motor learning in children with spina bifida: intact learning and performance on a ballistic task.

Learning and performance on a ballistic task were investigated in children with spina bifida meningomyelocele (SBM), with either upper level spinal lesions (n = 21) or lower level spinal lesions (n = 81), and in typically developing controls (n = 35). Participants completed three phases (20 trials each) of an elbow goniometer task that required a ballistic arm movement to move a cursor to one of two target positions on a screen, including (1) an initial learning phase, (2) an adaptation phase with a gain change such that recalibration of the ballistic arm movement was required, and (3) a learning reactivation phase under the original gain condition. Initial error rate, asymptotic error rate, and learning rate did not differ significantly between the SBM and control groups. Relative to controls, the SBM group had reduced volumes in the cerebellar hemispheres and pericallosal gray matter (the region including the basal ganglia), although only the pericallosal gray matter was significantly correlated with motor adaptation. Congenital cerebellar dysmorphology is associated with preserved motor skill learning on voluntary, nonreflexive tasks in children with SBM, in whom the relative roles of the cerebellum and basal ganglia may differ from those in the adult brain.

Adaptation, Physiological↗

Computational consequences of temporally asymmetric learning rules: I. Differential hebbian learning.

Temporally asymetric learning rules governing plastic changes in synaptic efficacy have recently been identified in physiological studies. In these rules, the exact timing of pre- and postsynaptic spikes is critical to the induced change of synaptic efficacy. The temporal learning rules treated in this article are approximately antisymmetric; the synaptic efficacy is enhanced if the postsynaptic spike follows the presynaptic spike by a few milliseconds, but the efficacy is depressed if the postsynaptic spike precedes the presynaptic spike. The learning dynamics of this rule are studied using a stochastic model neuron receiving a set of serially delayed inputs. The average change of synaptic efficacy due to the temporally antisymmetric learning rule is shown to yield differential Hebbian learning. These results are demonstrated with both mathematical analyses and computer simulations, and connections with theories of classical conditioning are discussed.

Animals↗

Comparison-based learning: effects of comparing instances during category learning.

When learning about a category, people often compare new instances with similar old instances and notice features common to the compared instances. Five experiments demonstrate that such comparisons cause features common to compared instances to be considered more important for the category than equally frequent features that are not common to compared instances. Experiment 1 shows that what is learned depends on which instances are compared. Experiment 2 investigates the conditions under which comparison-based learning occurs. The next experiments find that these comparisons affect subjective feature frequency (Experiment 3) and sensitivity to feature correlations (Experiment 4). Experiment 5 shows that comparisons during early learning affect what is learned from later instances. The discussion focuses on the implications for models of category representation.

Color Perception↗

Spatial attention and implicit sequence learning: evidence for independent learning of spatial and nonspatial sequences.

This research investigated whether regular spatial orienting sequences can be learned implicitly and independently of response requirements. In a new version of a serial response task introduced by M. J. Nissen and P. Bullemer (1987) participants had to discriminate between objects that could occur at different locations. Independent sequences determined the succession of locations and objects. Even participants who were not aware of any regularities exhibited evidence for learning of both sequences (Experiment 1). Experiment 2 showed that the joint learning of spatial and object sequences was as efficient as learning of single sequences and that it even occurred when learning required memory for past sequence elements and attention was blocked through a secondary tone-counting task. Results are consistent with the idea that independent systems may exist for the implicit acquisition of spatial and nonspatial regularities.

Adult↗

Why there are complementary learning systems in the hippocampus and neocortex: insights from the successes and failures of connectionist models of learning and memory.

Damage to the hippocampal system disrupts recent memory but leaves remote memory intact. The account presented here suggests that memories are first stored via synaptic changes in the hippocampal system, that these changes support reinstatement of recent memories in the neocortex, that neocortical synapses change a little on each reinstatement, and that remote memory is based on accumulated neocortical changes. Models that learn via changes to connections help explain this organization. These models discover the structure in ensembles of items if learning of each item is gradual and interleaved with learning about other items. This suggests that the neocortex learns slowly to discover the structure in ensembles of experiences. The hippocampal system permits rapid learning of new items without disrupting this structure, and reinstatement of new memories interleaves them with others to integrate them into structured neocortical memory systems.

Amnesia, Retrograde↗