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Formation of a motor memory by action observation.

Mirror neurons discharge with both action observation and action execution. It has been proposed that the mirror neuron system is instrumental in motor learning. The human primary motor cortex (M1) displays mirror activity in response to movement observation, is capable of forming motor memories, and is involved in motor learning. However, it is not known whether movement observation can lead directly to the formation of motor memories in the M1, which is considered a likely physiological step in motor learning. Here, we used transcranial magnetic stimulation (TMS) to show that observation of another individual performing simple repetitive thumb movements gives rise to a kinematically specific memory trace of the observed motions in M1. An extended period of observation of thumb movements that were oriented oppositely to the previously determined habitual directional bias increased the probability of TMS-evoked thumb movements to fall within the observed direction. Furthermore, the acceleration of TMS-evoked thumb movements along the principal movement axis and the balance of excitability of muscle representations active in the observed movements were altered in favor of the observed movement direction. These findings support a role for the mirror neuron system in memory formation and possibly human motor learning.

Adult↗

The role of knowledge of results frequency in learning through observation.

The authors examined whether reduced knowledge of results (KR) frequency during observation of a model's performance enhances learning. As they viewed a timing task, observers (n = 54) received KR about the model's performance on each trial (100% KR) or on 1 out of 3 trials (33% KR). Controls (n = 18) received only physical practice; they did not take part in the observation session. The authors also wanted to dissociate the guidance effect of KR during physical practice from the guidance role played by the representation acquired during observation. Therefore, following the observation phase, participants physically performed the task with either the same or a different KR frequency than that experienced during observation. The effects of observation and physical practice on learning were assessed in delayed retention tests. The beneficial effect of reduced KR frequency during observation continued for the following physical practice phases. Possible explanations as to why KR influences observational learning are discussed.

Adult↗

Involvement of the septohippocampal cholinergic system in representational memory.

To develop an animal model for testing muscarinic agonists, we examined the effects of cholinergic lesions with the ethylcholine aziridinium ion (AF64A) on two types of memory tasks. The tasks provided a distinction between representational and dispositional memory that could be measured in a single paradigm. Young, male Long-Evans rats were trained in a modified T-maze to learn both a discrimination task and a paired-run alternation task. Once animals learned the tasks, they were administered either saline or AF64A (5 nmol into each hippocampus) via stereotaxic technique. One week following surgery, saline-treated animals exhibited comparable performances (P greater than 0.2) on both the discrimination task (90.0 +/- 2.6% correct) and the alternation task (79.5 +/- 5.7%). In contrast, animals treated with AF64A showed a significant impairment of performance (P less than 0.005) on the alternation task (56.1 +/- 1.7%) as compared to the discrimination task (81.6 +/- 5.0%). Performance of the alternation task was significantly lower for AF64A-treated animals than for controls (P less than 0.02). AF64A-treated animals subsequently injected with pilocarpine (1.0 mg/kg, i.p.) showed moderate improvements in performance on the alternation task, while performance on the discrimination task remained unaffected. Immunocytochemical studies of choline acetyltransferase (ChAT) and tyrosine hydroxylase (TH) immunoreactivity indicated a loss of ChAT-positive cells in the septal region in AF64A-injected animals while TH-positive cells in the ventral tegmental area were unaffected by the treatment. The data suggest that AF64A can be used to produce selective lesions of the septohippocampal cholinergic system, which plays a greater role in representational memory than in dispositional memory.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Entorhinal cortex grid cells can map to hippocampal place cells by competitive learning.

'Grid cells' in the dorsocaudal medial entorhinal cortex (dMEC) are activated when a rat is located at any of the vertices of a grid of equilateral triangles covering the environment. dMEC grid cells have different frequencies and phase offsets. However, cells in the dentate gyrus (DG) and hippocampal area CA3 of the rodent typically display place fields, where individual cells are active over only a single portion of the space. In a model of the hippocampus, we have shown that the connectivity from the entorhinal cortex to the dentate granule cells could allow the dentate granule cells to operate as a competitive network to recode their inputs to produce sparse orthogonal representations, and this includes spatial pattern separation. In this paper we show that the same computational hypothesis can account for the mapping of EC grid cells to dentate place cells. We show that the learning in the competitive network is an important part of the way in which the mapping can be achieved. We further show that incorporation of a short term memory trace into the associative learning can help to produce the relatively broad place fields found in the hippocampus.

Action Potentials↗

Preserved learning in amnesic patients: perspectives from research on direct priming.

An important source of evidence for preserved learning ability in amnesic patients is provided by the phenomenon of repetition or direct priming: the facilitative effect of an encounter with an item on subsequent processing of the same item. In this article we consider a series of experiments that have attempted to clarify the nature of priming in amnesic patients and normal subjects. The experiments indicate that there may be two varieties of priming, one dependent upon the activation of preexisting memory representations, the other dependent upon newly formed memory representations. We suggest that the former type of priming is preserved even in severely amnesic patients, whereas the latter may be preserved only in mildly-to-moderately amnesic patients. Implications of research on preserved learning for the development of remedial strategies are also discussed.

Adult↗

Differential temporal evolution of post-training changes in regional brain glucose metabolism induced by repeated spatial discrimination training in mice: visualization of the memory consolidation process?

The present study analyses the effects of the stage of learning on the spatial patterns and time-course of [14C]glucose uptake in BALB/c mice brain regions produced by spatial discrimination training in an eight-arm radial maze. Our particular approach was designed to follow, during the post-training period, the level of functional activity in individual brain areas which may underlie the memory consolidation process. Regional mapping of relative [14C]glucose uptake was assessed at three post-training time intervals (5 min, 1 and 3 h) after either the first (Day 1), the fourth (Day 4) or the last (Day 9) daily training session of the discrimination task and compared with sham-conditioned animals placed in the same experimental environment. The results indicated that numerous subcortical and cortical brain regions exhibit metabolic alterations following the acquisition of the spatial discrimination task. These alterations, which were specifically related to learning since they did not appear in sham-conditioned animals, were functions both of the post-training interval studied and of the degree of mastery of the task. On Day 1, a progressive, time-dependent and sequential increase in labelling was found from subcortical (5 min post-training) to cortical regions (3 h post-training). On Day 4, a peak of cortical metabolic activation was identified at 1 h post-training. In contrast, on Day 9, maximum labelling was found 5 min post-training in all subcortical and cortical regions followed by a general monotonic decline at 1 and 3 h post-training. These findings, which show widely distributed changes of metabolic activity in the brain, are consistent with the hypothesis that learning involves distributed neural networks. The sequential activation from subcortical to cortical regions seems to indicate a general mechanism whose function would ultimately be to store cortical memory representations. The acquisition-dependent shifts in the patterns of post-training metabolic labelling observed as a function of task mastery may be taken to represent a visualization of the spatio-temporal evolution of the networks of brain structures actively engaged in the memory consolidation process. In particular, the present data suggest that the duration of post-acquisition memory processing is a function of the quantity of new information which has to be dealt with by the central nervous system.

Animals↗

Imagery of motor actions: differential effects of kinesthetic and visual-motor mode of imagery in single-trial EEG.

Single-trial motor imagery classification is an integral part of a number of brain-computer interface (BCI) systems. The possible significance of the kind of imagery, involving rather kinesthetic or visual representations of actions, was addressed using the following experimental conditions: kinesthetic motor imagery (MIK), visual-motor imagery (MIV), motor execution (ME) and observation of movement (OOM). Based on multi-channel EEG recordings in 14 right-handed participants, we applied a learning classifier, the distinction sensitive learning vector quantization (DSLVQ) to identify relevant features (i.e., frequency bands, electrode sites) for recognition of the respective mental states. For ME and OOM, the overall classification accuracies were about 80%. The rates obtained for MIK (67%) were better than the results of MIV (56%). Moreover, the focus of activity during kinesthetic imagery was found close to the sensorimotor hand area, whereas visual-motor imagery did not reveal a clear spatial pattern. Consequently, to improve motor-imagery-based BCI control, user training should emphasize kinesthetic experiences instead of visual representations of actions.

Adolescent↗

Explicit memory, repetition priming and cognitive skill learning in schizophrenia.

Explicit memory and two forms of implicit memory, repetition priming and cognitive skill learning, were examined in twenty-four schizophrenic patients and twenty-four normal control subjects previously matched for sex, age, and educational level. Two explicit tasks, free recall and frequency monitoring, and two implicit tasks, word completion and the Tower of Toronto puzzle, a variant of the Tower of Hanoi puzzle, were selected. The performance of schizophrenic patients was impaired in both explicit tasks, whereas repetition priming was intact; in the Tower of Toronto puzzle, a deficit was observed in problem solving, but not in skill learning. This dissociation between explicit and implicit memory is not entirely consistent with the hypothesis of a deficiency in effort-demanding information processing. It could be better accommodated by a model of a disturbance in the internal representation of context.

Adolescent↗

Importance of corpus callosum for visual receptive fields of single neurons in cat superior colliculus.

1. Section of the posterior two-thirds of the corpus callosum eliminates almost completely the response of superior colliculus (SC) neurons to stimulation of the contralateral eye in split-chiasm cats. On the contrary, the responsiveness of SC neurons to stimulation of the contralateral eye is not abolished by a transection of the posterior and tectal commissures leaving the corpus callosum intact. The callosal section also reduces the number of SC receptive fields abutting the vertical meridian in the ipsilateral eye of split-chiasm cats. 2. In cats with intact optic pathways, a similar callosal section abolishes the SC representation of the ipsilateral visual field in the ipsilateral eye and also reduces the number of receptive fields adjoining the vertical meridian in the same eye. In the contralateral eye, the SC representation of the ipsilateral visual field is reduced in extension to about one-fifth of that seen in cats with intact commissures. 3. The results suggest that the corpus callosum is the main pathway for cross-midline communication of visual information at not only the cortical, but also the midbrain level. The corpus callosum may subserve this function because it contains uninterrupted crossed corticotectal projections or because it transmits visual information from one hemisphere to contralateral cortical areas projecting ipsilaterally to SC. The latter hypothesis is more likely but, in any case, the findings imply that the lack of interhemispheric transfer of visual learning in cats with a chiasmatic and callosal section may depend on a midline disconnection of both subcortical and cortical visual centers. 4. The corpus callosum is also responsible for the representation of the ipsilateral visual field of the ipsilateral eye in the cat SC. The SC representation of the ipsilateral visual field in the contralateral eye is due, in minimal part, to direct retinotectal connections from temporal retina and, for the largest part, to the corpus callosum. 5. Finally, the corpus callosum contributes to the representation of the contralateral visual field near the vertical meridian of the temporal retina in both split-chiasm and normal cats. This is probably due to the scarcity of direct retinotectal projections from this part of the retina and to their supplementation by corticotectal neurons influenced by the callosal afferents.

Animals↗

Learning manual pursuit tracking skills in patients with Parkinson's disease.

Evidence from a number of sources identifies the putamen and its ultimate cortical projection sites as forming a possible substrate for motor learning. The present paper describes two experiments which explored motor learning of a pursuit tracking task under first (position) and second (velocity) order control dynamics, in patients with Parkinson's disease on and off (experiment 2 only) their normal dopaminergic medication. In neither experiment did the medicated patients show evidence of significant impairment in learning the tasks. In the velocity tracking task, however, the patients off medication showed significantly less improvement in performance with practice. The discussion considers a number of possible interpretations of this finding. Contemporary cognitive theories of motor learning consider behavioural change with practice to be the combined action of an automatic procedural system, together with input from a conscious declarative system. Development of declarative knowledge about the task may have changed the nature of the process involved, from a visually guided task to a more predictive one based upon an internal representation. Evidence from various sources suggests that patients with Parkinson's disease have particular problems with this mode of control, thus making the task more difficult. It is suggested that motor control deficits have not been adequately considered in previous studies on motor learning, and that the evidence from clinical studies for a role of the putamen/supplementay motor area in motor learning remains equivocal.

Aged↗

Complexity and non-commutativity of learning operations on graphs.

We present results from numerical studies of supervised learning operations in small recurrent networks considered as graphs, leading from a given set of input conditions to predetermined outputs. Graphs that have optimized their output for particular inputs with respect to predetermined outputs are asymptotically stable and can be characterized by attractors, which form a representation space for an associative multiplicative structure of input operations. As the mapping from a series of inputs onto a series of such attractors generally depends on the sequence of inputs, this structure is generally non-commutative. Moreover, the size of the set of attractors, indicating the complexity of learning, is found to behave non-monotonically as learning proceeds. A tentative relation between this complexity and the notion of pragmatic information is indicated.

Artificial Intelligence↗

A configural analysis for feature-negative discrimination learning.

In 4 experiments, rats were initially trained with an A+ AXo discrimination in which Stimulus A by itself signaled the delivery of food, A+, whereas the simultaneous presentation of A and X was followed by nothing, AXo. In each experiment X was then paired with food prior to a test phase in which A and X were again presented for a discrimination. The discrimination was of the form A+ AXo in Experiments 1 and 2, whereas it was of the form X+ AXo for Experiments 3 and 4. In all 4 experiments the test discrimination was acquired more rapidly than a control discrimination. The results are interpreted in terms of the original A+ AXo discrimination resulting in the growth of an association between a representation of the entire AX compound and the effects of nonreinforcement.

Animals↗

The bilingual brain: cerebral representation of languages.

The present article deals with theoretical and experimental aspects of language representation in the multilingual brain. Two general approaches were adopted in the study of the bilingual brain. The study of bilingual aphasics allows us to describe dissociations and double dissociations between the different subcomponents of the various languages. Furthermore, symptoms peculiar to bilingual aphasia were reported (pathological mixing and switching and translations disorders) which allowed the correlation of some abilities specific to bilinguals with particular neurofunctional systems. Another approach to the study of the bilingual brain is of the experimental type, such as electrophysiological investigations (electrocorticostimulation during brain surgery and event-related potentials) and functional neuroanatomy studies (positron emission tomography and functional magnetic resonance imaging). Functional neuroanatomy studies investigated the brain representation of languages when processing lexical and syntactic stimuli and short stories. Neurophysiologic and neuroimaging studies evidenced a similar cerebral representation of L1 and L2 lexicons both in early and late bilinguals. The representation of grammatical aspects of languages seems to be different between the two languages if L2 is acquired after the age of 7, with automatic processes and correctness being lower than those of the native language. These results are in line with a greater representation of the two lexicons in the declarative memory systems, whereas morphosyntactic aspects may be organized in different systems according to the acquisition vs learning modality.

Brain↗

Post-project appraisals in adaptive management of river channel restoration.

Post-project appraisals (PPAs) can evaluate river restoration schemes in relation to their compliance with design, their short-term performance attainment, and their longer-term geomorphological compatibility with the catchment hydrology and sediment transport processes. PPAs provide the basis for communicating the results of one restoration scheme to another, thereby improving future restoration designs. They also supply essential performance feedback needed for adaptive management, in which management actions are treated as experiments. PPAs allow river restoration success to be defined both in terms of the scheme attaining its performance objectives and in providing a significant learning experience. Different levels of investment in PPA, in terms of pre-project data and follow-up information, bring with them different degrees of understanding and tbus different abilities to gauge both types of success. We present four case studies to illustrate how the commitment to PPA has determined the understanding achieved in each case. In Moore's Gulch (California, USA), understanding was severely constrained by the lack of pre-project data and post-implementation monitoring. Pre-project data existed for the Kitswell Brook (Hertfordshire, UK), but the monitoring consisted only of one site visit and thus the understanding achieved is related primarily to design compliance issues. The monitoring undertaken for Deep Run (Maryland, USA) and the River Idle (Nottinghamshire, UK) enabled some understanding of the short-term performance of each scheme. The transferable understanding gained from each case study is used to develop an illustrative five-fold classification of geomorphological PPAs (full, medium-term, short-term, one-shot, and remains) according to their potential as learning experiences. The learning experience is central to adaptive management but rarely articulated in the literature. Here, we gauge the potential via superimposition onto a previous schematic representation of the adaptive management process by Haney and Power (1996). Using PPAs wisely can lead to cutting-edge, complex solutions to river restoration challenges.

Communication↗

Priming contour-deleted images: evidence for intermediate representations in visual object recognition.

The speed and accuracy of perceptual recognition of a briefly presented picture of an object is facilitated by its prior presentation. Picture priming tasks were used to assess whether the facilitation is a function of the repetition of: (a) the object's image features (viz., vertices and edges), (b) the object model (e.g., that it is a grand piano), or (c) a representation intermediate between (a) and (b) consisting of convex or singly concave components of the object, roughly corresponding to the object's parts. Subjects viewed pictures with half their contour removed by deleting either (a) every other image feature from each part, or (b) half the components. On a second (primed) block of trials, subjects saw: (a) the identical image that they viewed on the first block, (b) the complement which had the missing contours, or (c) a same name-different exemplar of the object class (e.g., a grand piano when an upright piano had been shown on the first block). With deletion of features, speed and accuracy of naming identical and complementary images were equivalent, indicating that none of the priming could be attributed to the features actually present in the image. Performance with both types of image enjoyed an advantage over that with the different exemplars, establishing that the priming was visual rather than verbal or conceptual. With deletion of the components, performance with identical images was much better than that with their complements. The latter were equivalent to the different exemplars, indicating that all the visual priming of an image of an object is through the activation of a representation of its components in specified relations. In terms of a recent neural net implementation of object recognition (Hummel & Biederman, in press), the results suggest that the locus of object priming may be at changes in the weight matrix for a geon assembly layer, where units have self-organized to represent combinations of convex or singly concave components (or geons) and their attributes (e.g., aspect ratio, orientation, and relations with other geons such as TOP-OF). The results of these experiments provide evidence for the psychological reality of intermediate representations in real-time visual object recognition.

Adult↗

The perception of identity by 6 1/2-month-old infants.

In Study 1, sixteen 6 1/2-month-olds were habituated to a Reversible stimulus (an upright face that could be perceived as an entirely different upright face when it was rotated 180 degrees) and to a Nonreversible stimulus (a face that could be perceived as upright in only one orientation). Following habituation for each type of stimulus, test trials paired the habituated face with a novel stimulus (an inversion of the same face). For both Reversible and Nonreversible stimuli, the physical difference between the old and new test stimuli was the same (a 180 degrees rotation); however, infants devoted more visual attention to the 180 degrees rotation only when it was a Reversible face, suggesting that the identity change was detected. Experiment 2 ruled out the explanation that infants might have failed to dishabituate to the inversion of the Nonreversible stimulus because they could not remember it. Results are interpreted as evidence that 6 1/2-month-old infants are not limited to face recognition based on similarity in pattern arrangement alone, but are capable of processing faces at a representational level.

Attention↗

On the nature of animal thinking.

Recent attempts to teach apes rudimentary grammatical skills have produced negative results. The basic obstacle appears to be at the level of the individual symbol which, for apes, functions only as a demand. Little evidence exists that apes can use symbols as names, that is, as a means of simply transmitting information. Even though non-human animals lack linguistic competence, much evidence has accumulated recently that a variety of animals can form representations of particular features of their environment. What then is the non-verbal nature of animal representations? This question will be discussed with reference to recent investigations of animal memory--in particular, studies concerned with a pigeon's ability to learn "lists" of arbitrary elements. While learning to produce a particular sequence of four elements (colors), pigeons also acquire knowledge about the relation between non-adjacent elements and about the ordinal position of a particular element. Learning to produce a particular sequence also facilitates the discrimination of that sequence from other sequences.

Animals↗

How chicks make memories: the cellular cascade from c-fos to dendritic remodelling.

Training chicks on a one-trial passive avoidance task results in a cellular cascade over the subsequent hours. Phosphorylation of the presynaptic phosphokinase C substrate B-50 is followed by immediate-early gene expression and increased synthesis of pre- and postsynaptic glycoproteins, increases in dendritic spine densities, synapse and synaptic vesicle numbers, and a prolonged increase in neuronal bursting. Many of these effects have been localized to two forebrain regions: the left intermediate medial hyperstriatum ventrale and the lobus parolfactorius. Pretraining lesions in the left intermediate medial hyperstriatum ventrale, or post-training lesions in the lobus parolfactorius result in amnesia. These and related results lead to models of memory storage based on multiple representation by way of synaptic stabilization through glycoprotein synaptic recognition molecules.

Animals↗