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PubMed2004

Infant grasp learning: a computational model.

This paper presents ILGM (the Infant Learning to Grasp Model), the first computational model of infant grasp learning that is constrained by the infant motor development literature. By grasp learning we mean learning how to make motor plans in response to sensory stimuli such that open-loop execution of the plan leads to a successful grasp. The open-loop assumption is justified by the behavioral evidence that early grasping is based on open-loop control rather than on-line visual feedback. Key elements of the infancy period, namely elementary motor schemas, the exploratory nature of infant motor interaction, and inherent motor variability are captured in the model. In particular we show, through computational modeling, how an existing behavior (reaching) yields a more complex behavior (grasping) through interactive goal-directed trial and error learning. Our study focuses on how the infant learns to generate grasps that match the affordances presented by objects in the environment. ILGM was designed to learn execution parameters for controlling the hand movement as well as for modulating the reach to provide a successful grasp matching the target object affordance. Moreover, ILGM produces testable predictions regarding infant motor learning processes and poses new questions to experimentalists.

Computational Biology↗
PubMed2002

Interindividual variability of learning in stereoacuity.

BACKGROUND: In the evaluation of therapies aiming at binocular vision, for instance by the use of prisms or orthoptic training in the case of heterophoria, stereoacuity is often the primary outcome measure. To assess therapeutic effects it is necessary to separate them from perceptual learning with repeated testing. Learning stereoacuity has been investigated only in a few studies with up to six subjects. METHODS: To ascertain the interindividual variability of learning in stereoacuity we examined 24 subjects, 12 with and 12 without experience in psychophysical experiments. In a two-alternative forced-choice paradigm, subjects reported whether a vertical bar appeared in front of or behind a reference frame. Estimates of stereo threshold were obtained using an adaptive staircase procedure ("best PEST"). RESULTS: We found a highly significant learning effect ( P<0.0001) with a marked interindividual variability. In some subjects the stereoacuity improved by a factor of >30 and in others it did not improve at all. The median of the learning factor was 1.7. There was no significant difference between novices and experienced subjects. CONCLUSION: The great interindividual variability of learning in stereoacuity has important implications for therapeutic tests that use stereoacuity as an outcome measure: To distinguish therapeutic effects from improvements due to repeated testing, each subject's individual learning behaviour has to be taken into account, for example by starting out with an adequate training phase. The number of test repetitions required to reach a fairly constant level appears to be similar among individuals: in our paradigm most of the learning occurred within the first six blocks with 100 target presentations each.

Adult↗
PubMed2001

Statistical characteristics of climbing fiber spikes necessary for efficient cerebellar learning.

Mean firing rates (MFRs), with analogue values, have thus far been used as information carriers of neurons in most brain theories of learning. However, the neurons transmit the signal by spikes, which are discrete events. The climbing fibers (CFs), which are known to be essential for cerebellar motor learning, fire at the ultra-low firing rates (around 1 Hz), and it is not yet understood theoretically how high-frequency information can be conveyed and how learning of smooth and fast movements can be achieved. Here we address whether cerebellar learning can be achieved by CF spikes instead of conventional MFR in an eye movement task, such as the ocular following response (OFR), and an arm movement task. There are two major afferents into cerebellar Purkinje cells: parallel fiber (PF) and CF, and the synaptic weights between PFs and Purkinje cells have been shown to be modulated by the stimulation of both types of fiber. The modulation of the synaptic weights is regulated by the cerebellar synaptic plasticity. In this study we simulated cerebellar learning using CF signals as spikes instead of conventional MFR. To generate the spikes we used the following four spike generation models: (1) a Poisson model in which the spike interval probability follows a Poisson distribution, (2) a gamma model in which the spike interval probability follows the gamma distribution, (3) a max model in which a spike is generated when a synaptic input reaches maximum, and (4) a threshold model in which a spike is generated when the input crosses a certain small threshold. We found that, in an OFR task with a constant visual velocity, learning was successful with stochastic models, such as Poisson and gamma models, but not in the deterministic models, such as max and threshold models. In an OFR with a stepwise velocity change and an arm movement task, learning could be achieved only in the Poisson model. In addition, for efficient cerebellar learning, the distribution of CF spike-occurrence time after stimulus onset must capture at least the first, second and third moments of the temporal distribution of error signals.

Action Potentials↗
PubMed2002

Learning to discriminate simple sinusoidal gratings is task specific.

Learning transfer effects of a modified spatial frequency discrimination task with simple sinusoidal gratings on various untrained test tasks, testing performance in relative position, local width and global size discrimination, have been investigated. Six subjects were exposed to grating stimuli of varying spatial frequency and size but constant relative position while they had to respond only to spatial frequency. In the course of 7 consecutive days all subjects showed significant reduction of spatial frequency discrimination thresholds. Comparison of discrimination thresholds for the untrained test tasks, taken before and after the learning epoch, reveals complete learning transfer to spatial frequency discrimination with gratings of constant size and variable relative position, but complete lack of transfer when grating spatial frequency is shifted about one octave lower. Further, there is improvement of relative position discrimination and width discrimination of single luminance bars, which is not very specific for the learnt spatial frequency. Although size variation was part of the learning procedure, discrimination of global stimulus size did not improve. Generally, the observed scheme of learning transfer reveals that there is learning only for stimulus attributes that are behaviorally relevant in the learning task. The differential scheme of improvement and code usage in the test tasks strongly indicates involvement of higher stages capable of independent access to different coding domains, as well as attentionally guided attribute selection and suppression. Supported by other recent findings, it is suggested that discrimination learning can be understood as a higher level process of gradual refinement of code selection out of a rich code base provided by lower level stages.

Adult↗
PubMed2006

Learning by the parasitoid wasp, Aphidius ervi (Hymenoptera: Braconidae), alters individual fixed preferences for pea aphid color morphs.

Learning, defined as changes in behavior that occur due to past experience, has been well documented for nearly 20 species of hymenopterous parasitoids. Few studies, however, have explored the influence of learning on population-level patterns of host use by parasitoids in field populations. Our study explores learning in the parasitoid Aphidius ervi Haliday that attacks pea aphids, Acyrthosiphon pisum (Harris). We used a sequence of laboratory experiments to investigate whether there is a learned component in the selection of red or green aphid color morphs. We then used the results of these experiments to parameterize a model that examines whether learned behaviors can explain the changes in the rates of parasitism observed in field populations in South-central Wisconsin, USA. In the first of two experiments, we measured the sequence of host choice by A. ervi on pea aphid color morphs and analyzed this sequence for patterns in biased host selection. Parasitoids exhibited an inherent preference for green aphid morphs, but this preference was malleable; initial encounters with red aphids led to a greater chance of subsequent orientation towards red aphids than predicted by chance. In a second experiment, we found no evidence that parasitoids specialize on red or green morphs; for the same parasitoids tested in trials separated by 2 h, color preference in the first trial did not predict color preference in the second, as would be expected if they differed in fixed preferences or exhibited long-term (> 2 h) learning. Using data from the two experiments, we parameterized a population dynamics model and found that learning of the magnitude observed in our experiments leads to biased parasitism towards the most common color morph. This bias is sufficient to explain changes in the ratio of aphid color morphs observed in field sites over multiple years. Our study suggests that for even relatively simple organisms, learned behaviors may be important for explaining the population dynamics of their hosts.

Animals↗
PubMed2006

Orientation in the cuttlefish Sepia officinalis: response versus place learning.

Several studies have demonstrated that mammals, birds and fish use comparable spatial learning strategies. Unfortunately, except in insects, few studies have investigated spatial learning mechanisms in invertebrates. Our study aimed to identify the strategies used by cuttlefish (Sepia officinalis) to solve a spatial task commonly used with vertebrates. A new spatial learning procedure using a T-maze was designed. In this maze, the cuttlefish learned how to enter a dark and sandy compartment. A preliminary test confirmed that individual cuttlefish showed an untrained side-turning preference (preference for turning right or left) in the T-maze. This preference could be reliably detected in a single probe trial. In the following two experiments, each individual was trained to enter the compartment opposite to its side-turning preference. In Experiment 1, distal visual cues were provided around the maze. In Experiment 2, the T-maze was surrounded by curtains and two proximal visual cues were provided above the apparatus. In both experiments, after acquisition, strategies used by cuttlefish to orient in the T-maze were tested by creating a conflict between the formerly rewarded algorithmic behaviour (turn, response learning) and the visual cues identifying the goal (place learning). Most cuttlefish relied on response learning in Experiment 1; the two strategies were used equally often in Experiment 2. In these experiments, the salience of cues provided during the experiment determined whether cuttlefish used response or place learning to solve this spatial task. Our study demonstrates for the first time the presence of multiple spatial strategies in cuttlefish that appear to closely parallel those described in vertebrates.

Animals↗
PubMed2006

Influence of learning resources on study time and achievement scores in a problem-based curriculum.

Many aspects of the problem-based learning cycle aim at stimulating students to elaborate on their knowledge. This elaborative process is considered to be one of the most effective study aids for learning and applying information. One of the opportunities to stimulate this elaboration process is by offering students more, and more diverse, learning resources. Under laboratory conditions, increasing the richness and diversity of the learning materials has often been shown to support students in learning, remembering and applying information. In this study, a curricular perspective was adopted on the question whether students would benefit from more learning resources. Mean study times and achievement scores for 28 problem-based courses were calculated and included in the analyses. Independent variables in both analyses were the number of primary learning resources (i.e., resources students were recommended to study) and the number of supplementary resources (i.e., related, but non-essential resources). Results showed a significant increase in achievement scores for courses with more primary learning resources, but the effect of supplementary resources was limited.

Educational Measurement↗
PubMed1993

Physical activity enhances spatial learning performance with an associated alteration in hippocampal protein kinase C activity in C57BL/6 and DBA/2 mice.

The effects of physical activity on spatial learning performance and associated hippocampal functioning were examined in C57BL/6Ibg (C57) and DBA/2Ibg (DBA) mice. C57 and DBA mice, 3 months of age, were subjected to 8 weeks of a physical activity regime (consisting of moderate-pace treadmill running 5 days/week, 60 min/day, 0% grade, 12 m/min) or remained sedentary in their cages. Mice were then tested on the Morris water maze task for 6 days followed by 12 days of testing on the place learning-set task (8 trials/day with each task). Both C57 and DBA run mice showed no difference in swim speed compared to controls. Hippocampal protein kinase C (PKC) activity was measured in cytosolic, loosely bound, and membrane-bound homogenate fractions. Mice subjected to the physical activity protocol were compared to sedentary controls from the same set of litters. Physical activity produced a 2- to 12-fold enhancement in spatial learning performance on both the Morris (P < 0.0001) and place learning-set (P < 0.02) probe trials in both C57 and DBA mice. DBA mice, which characteristically perform poorly in comparison to C57 mice, were enhanced to perform similarly to C57 control mice. This physical activity-induced enhancement in spatial learning performance was accompanied by alterations in hippocampal bound PKC activity (P < 0.05). These data provide further support for our previous hypotheses of a PKC activity involvement in spatial learning and enhancement of spatial learning performance in rodents by physical activity. In addition, these data indicate that hippocampal PKC activity may be involved in the physical activity-induced enhancement of spatial learning performance.

Analysis of Variance↗
PubMed1991

U-shaped learning and frequency effects in a multi-layered perceptron: implications for child language acquisition.

A three-layer back-propagation network is used to implement a pattern association task in which four types of mapping are learned. These mappings, which are considered analogous to those which characterize the relationship between the stem and past tense forms of English verbs, include arbitrary mappings, identity mappings, vowel changes, and additions of a suffix. The degree of correspondence between parallel distributed processing (PDP) models which learn mappings of this sort (e.g., Rumelhart & McClelland, 1986, 1987) and children's acquisition of inflectional morphology has recently been at issue in discussions of the applicability of PDP models to the study of human cognition and language (Pinker & Mehler, 1989; Bever, in press). In this paper, we explore the capacity of a network to learn these types of mappings, focusing on three major issues. First, we compare the performance of a single-layered perceptron similar to the one used by Rumelhart and McClelland with a multi-layered perceptron. The results suggest that it is unlikely that a single-layered perceptron is capable of finding an adequate solution to the problem of mapping stems and past tense forms in input configurations that are sufficiently analogous to English. Second, we explore the input conditions which determine learning in these networks. Several factors that characterize linguistic input are investigated: (a) the nature of the mapping performed by the network (arbitrary, suffixation, identity, and vowel change); (b) the competition effects that arise when the task demands simultaneous learning of distinct mapping types; (c) the role of the type and token frequency of verb stems; and (d) the influence of phonological subregularities in the irregular verbs. Each of these factors is shown to have selective consequences on both successful and erroneous performance in the network. Third, we outline several types of systems which could result in U-shaped acquisition, and discuss the ways in which learning in multi-layered networks can be seen to capture several characteristics of U-shaped learning in children. In general, these models provide information about the role of input in determining the kinds of errors that a network will produce, including the conditions under which rule-like behavior and U-shaped learning will and will not emerge. The results from all simulations are discussed in light of behavioral data on children's acquisition of the past tense and the validity of drawing conclusions about the acquisition of language from models of this sort.

Child, Preschool↗
PubMed1994

Altered serial position learning after frontal lobe lesion.

The serial position function is a powerful and highly reliable feature of human learning, with well-described primacy and recency effects. We tested the hypothesis that frontal lobe lesions in patients would disrupt the serial position function since such patients are known to have disturbed temporal ordering, learning in the presence of interference, encoding and organizational approaches to learning. Performance was compared in patients with focal, acquired lesions of frontal and non-frontal cortices, using a standardized paradigm of verbal list learning. Results indicated a similar pattern of performance on first trial learning for the two groups. However, across learning trials, frontal lesion subjects failed to maintain significant primacy and recency effects. Non-frontal lesion subjects consistently showed the expected U-shaped serial position curve across all trials. Subjective organization in learning was particularly deficient in the dorsolateral frontal lesion subjects. We propose that serial position effects are qualitatively different after frontal lobe lesion, being transitory and prone to alteration by the cumulative effects of disturbed temporal-spatial processing across learning trials.

Adult↗
PubMed1994

When implicit learning fails: amnesia and the problem of error elimination.

We propose that one of the major functions of explicit memory is the elimination of learning errors. The hypothesis is explored by means of a stem completion task in which subjects are presented with stems having many potential completions, and in the initial phase are either encouraged to guess, the "errorful" procedure, or are provided with the correct completion, the "errorless" condition. Learning is then tested over a sequence of nine trials. The performance of amnesic subjects who are assumed to have good implicit but bad explicit learning is compared with that of normal elderly subjects, who are assumed to have an intermediate level of explicit learning skill, and young controls who are expected to be high in both implicit and explicit learning capacity. As predicted, errorless learning is beneficial, with the effect being particularly marked for the amnesic group. A detailed analysis of the intrusion errors supports an interpretation of the results in terms of the relative contribution to the three groups of implicit learning, which is assumed to be particularly vulnerable to interference. Implications for the analysis of normal learning, and for the rehabilitation of brain damaged patients are discussed.

Adult↗
PubMed1985

Involvement of the anterior insular gustatory neocortex in taste-potentiated odor aversion learning.

When an odor conditioned stimulus (CS) precedes illness (unconditioned stimulus; UCS), rats acquire relatively weak odor aversions. Conversely, when a compound odor-taste (flavor) CS precedes illness, rats acquire robust aversions both to the odor and to the taste components of a compound flavor CS. Thus, tastes potentiate odor-illness aversions during toxiphobic conditioning. Such conditioning effects have been referred to as taste-potentiated odor aversion learning (POA). Previous neurobehavioral experiments have shown that the anterior insular gustatory neocortex contributes to conditioned taste aversion (CTA) learning. The present experiment examined the involvement of the anterior insular gustatory neocortex in CTA learning and POA learning. To that end, four distinct groups of rats received bilateral electrolytic lesion placements in the orbitofrontal neocortex, the "somatic" gustatory neocortex, the anterior insular gustatory neocortex or the posterior insular neocortex. Control animals received anesthesia only. Subgroups of animals thereafter received aversion conditioning using either an odor (almond) CS or a compound odor-taste (almond-saccharin) CS. Aversions to the almond odorant and/or saccharin tastant were evaluated during extinction. Results indicated that animals lacking orbitofrontal neocortex or posterior insular neocortex acquired normal CTAs and POAs. Animals lacking somatic gustatory neocortex exhibited impaired CTA learning, yet those animals showed normal POA learning. Lesions centered in the anterior insular neocortex impaired both CTA learning and POA learning. These results demonstrate that the insular gustatory neocortex is uniquely involved in the higher-order integration of odors, tastes and illness.

Animals↗
PubMed1985

A comparison of shock avoidance learning in headless cockroaches in leg-lifting and lowering task.

Headless cockroaches were trained either to lift or to lower the prothoracic leg to avoid electric shocks. The learning process in both cases was very distinct. However, the learning performance in the lifting was better than in the lowering task. The reason for this poorer learning in the lowering task is due to the unconditioned leg raising movement of the animals to electric shocks. This unconditioned response affects the headless cockroaches avoid shocks in the lifting task by escape learning, whereas they avoid shocks in the lowering task by true avoidance learning. Injection of 30 microliter of Ringer's solution has no significant effect on the learning process in the lifting task. In the lowering task, however, Ringer's solution impairs the learning ability considerably. Crude extract of corpora cardiaca strongly inhibits the learning of both tasks.

Animals↗
PubMed1993

Salivary testosterone in children with and without learning disabilities.

Previous studies have indicated that the sex steroids have organizational effects upon neural tissue and that abnormal secretion during development may lead to functional anomalies. In this study, we explore the possibility of prepubertal steroid hormone involvement in the etiology of learning disabilities. Salivary testosterone levels in 264 children without learning disabilities (133 males, 131 females) were measured and compared to that in 32 children with learning disabilities (25 males, 7 females). The presence of learning disabilities was significantly associated with higher salivary testosterone. Data from equivalent samples of learning-disabled and control subjects also were compared separately because of disparities in sample size and variable distribution in the total group analysis. A 32-member sample of nonlearning-disabled children was created by randomly selecting individuals who exactly matched the age, race, and sex characteristics of the learning-disabled group. The matched analysis further substantiated the association between testosterone secretion and learning disabilities. Thus, it is possible that some learning disabilities may be associated in part with abnormal testosterone levels.

Adolescent↗
PubMed1994

LTP of mossy fiber-stimulated potentials in CA3 during learning in rats.

The study was done to determine whether long-term potentiation (LTP) of the potentials occurs at mossy fiber (MF)-CA3 synapses with the advance of learning in unrestrained and unanesthetized rats. The rats were divided into two groups, the test and control groups. The test group was given daily learning tasks in the radial arm maze, whereas the control group was similarly handled without learning tasks. Complete acquisition of learning was observed in the test group on day 5, and the learning was maintained over 3 days. Under freely moving conditions, a significant increase in population spikes (PS) elicited by MF stimulation with the progress in learning was observed in the test group, and the PS potentiation remained stable after day 4. Furthermore, on day 7, when MF stimulation-induced PS in the test group were compared with that in the control group in pentobarbital-anesthetized rats, the responses were comparatively higher in the former. As the training-induced PS potentiation in CA3 occurred with the advance of learning, these findings suggest that LTP in CA3 induced by learning may be related to memory storage.

Animals↗
PubMed1991

Effects of ketamine and 1-glutamic acid diethyl ester on concept learning in rats.

The effects of ketamine, an NMDA receptor antagonist, and 1-glutamic acid diethyl ester (LGDE), a non-NMDA glutamate antagonist, were evaluated in the acquisition of concept learning in a water maze. In concept learning, the rats must locate an invisible platform whose location changes from day to day. In spatial learning (Morris task), the rats must locate an invisible (or visible) platform whose location does not change. Ketamine increased quadrant entries at 5, 10 and 20 mg/kg, and latencies at 10 and 20 mg/kg on the final two days of training on the concept task. At 5 mg/kg ketamine disrupted concept learning but not spatial learning or visuo-motor coordination as assessed by invisible and visible platform conditions of the Morris maze. Progressively higher doses of ketamine affected first the invisible condition and then the visible platform condition. On the other hand, LGDE did not affect the Morris task at any dose. However, there was no decrease in latencies over days in concept learning at the two highest doses (240 and 360 mg/kg) of LGDE. Thus LGDE appeared to slow down decision time in the concept task but not the spatial task in the absence of an effect on quadrant entries in any version. These results indicate that NMDA receptors are involved in spatial and concept learning. Non-NMDA receptors appear to be involved only in concept learning.

Animals↗
PubMed1984

Decortication abolishes place but not cue learning in rats.

The experiments examined whether decorticate rats are able to acquire a place learning strategy, as compared with a cue learning strategy, to successfully navigate from one place to another and whether the hippocampus, in the absence of the neocortex, contributes to successful performance. Decorticate rats, with or without hippocampectomy, were unable to locate an "invisible" platform submerged at a fixed place in a tank of cool water (made opaque by milk), rather they scrabbled at the edges of the tank and failed even to initiate search strategies. They were able to learn to swim directly to the platform if it was visible. Their ability to find the hidden platform was not enhanced by presurgical experience or two-stage ablations with training before and after ablations. When pretrained on the cue task and tested on the place task, they learned to inhibit scrabbling at the tank edges and "search" in a haphazard fashion for the hidden platform, but they never learned to swim directly toward it. When decorticate rats, trained on the cue task, received superior colliculus or basal ganglia removal in a second operation, cue learning was abolished. Hippocampal removal after decortication left performance on the cue task unaffected. The results demonstrate: (1) the integrity of the neocortex is essential for place learning; (2) the brainstem, including superior colliculus and basal ganglia, is sufficient for cue learning; and (3) in the absence of the neocortex the hippocampus plays no role in guiding either type of navigation. It is concluded that sensorimotor subsystems of the forebrain play a special role as detector-response systems for guiding behaviour in response to constellations of distal stimuli, whereas subcortical structures are sufficient for navigation to a single stimulus.

Animals↗
PubMed1988

Inferotemporal-frontal disconnection and fornix transection in visuomotor conditional learning by monkeys.

The first 2 experiments examined the ability of monkeys (Macaca fascicularis) to learn a series of visuomotor conditional discrimination problems for food reward. In each discrimination problem there were 2 visual stimuli, which were different from those in any previous discrimination problem. Each trial within a problem presented either one visual stimulus or the other, and the stimulus indicated which of 2 motor responses, tap or hold, was correct; the motor responses were defined in such a way as to be mutually exclusive. Successive problems were each trained to a criterion of correct performance. In the first experiment, it was found that fornix transection did not impair monkey's learning rate in this task. This result contrasts with previous results showing an impairment of learning rate following fornix transection in visuo-motor conditional discriminations in which the 2 motor responses were differentiated from each other by their spatial position or direction. The present result shows that the earlier demonstrated impairments are specific to spatial responses. In the second experiment, learning rate in the present task was found to be impaired by the combination of a unilateral inferotemporal ablation, contralateral to the hand used in the task by the animal, with a transection of the anterior corpus callosum. This result suggests that cortico-cortical interaction between the inferotemporal area and the frontal lobe contralateral to the hand in use is necessary for efficient learning in this task. The third experiment examined simultaneous 2-choice visual discrimination learning in the animals which had participated in experiment 2. In contrast to the results of Expt. 2, learning rate in this task was unimpaired. Experiment 3 shows that the impairment observed in Expt. 2 is not a general impairment of visual learning.

Animals↗