[Selectivity in stimulus and memory representation].
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Adult↗
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Representation for discovery of protein motifs.
There are several dimensions and levels of complexity in which information on protein motifs may be available. For example, one-dimensional sequence motifs may be associated with secondary structure identifiers. Alternatively, three-dimensional information on polypeptide segments may be used to induce prototypical three-dimensional structure templates. This paper surveys various representations encountered in the protein motif discovery literature. Many of the representations are based on incompatible semantics, making difficult the comparison and combination of previous results. To make better use of machine learning techniques and to provide for an integrated knowledge representation framework, a general representation language--in which all types of motifs can be encoded and given a uniform semantics--is required. In this paper we propose such a model, called a spatial description logic, and present a machine learning approach based on the model.
Amino Acids↗
Neuronal substrates of motor learning in the velocity storage generated during optokinetic stimulation in the squirrel monkey.
Chronic motor learning in the vestibuloocular reflex (VOR) results in changes in the gain of this reflex and in other eye movements intimately associated with VOR behavior, e.g., the velocity storage generated by optokinetic stimulation (OKN velocity storage). The aim of the present study was to identify the plastic sites responsible for the change in OKN velocity storage after chronic VOR motor learning. We studied the neuronal responses of vertical eye movement flocculus target neurons (FTNs) during the optokinetic after-nystagmus (OKAN) phase of the optokinetic response (OKR) before and after VOR motor learning. Our findings can be summarized as follows. 1) Chronic VOR motor learning changes the horizontal OKN velocity storage in parallel with changes in VOR gain, whereas the vertical OKN velocity storage is more complex, increasing with VOR gain increases, but not changing following VOR gain decreases. 2) FTNs contain an OKAN signal having opposite directional preferences after chronic high versus low gain learning, suggesting a change in the OKN velocity storage representation of FTNs. 3) Changes in the eye-velocity sensitivity of FTNs during OKAN are correlated with changes in the brain stem head-velocity sensitivity of the same neurons. And 4) these changes in eye-velocity sensitivity of FTNs during OKAN support the new behavior after high gain but not low gain learning. Thus we hypothesize that the changes observed in the OKN velocity storage behavior after chronic learning result from changes in brain stem pathways carrying head velocity and OKN velocity storage information, and that a parallel pathway to vertical FTNs changes its OKN velocity storage representation following low, but not high, gain VOR motor learning.
Adaptation, Physiological↗
Formation of temporal-feature maps by axonal propagation of synaptic learning.
Computational maps are of central importance to a neuronal representation of the outside world. In a map, neighboring neurons respond to similar sensory features. A well studied example is the computational map of interaural time differences (ITDs), which is essential to sound localization in a variety of species and allows resolution of ITDs of the order of 10 micros. Nevertheless, it is unclear how such an orderly representation of temporal features arises. We address this problem by modeling the ontogenetic development of an ITD map in the laminar nucleus of the barn owl. We show how the owl's ITD map can emerge from a combined action of homosynaptic spike-based Hebbian learning and its propagation along the presynaptic axon. In spike-based Hebbian learning, synaptic strengths are modified according to the timing of pre- and postsynaptic action potentials. In unspecific axonal learning, a synapse's modification gives rise to a factor that propagates along the presynaptic axon and affects the properties of synapses at neighboring neurons. Our results indicate that both Hebbian learning and its presynaptic propagation are necessary for map formation in the laminar nucleus, but the latter can be orders of magnitude weaker than the former. We argue that the algorithm is important for the formation of computational maps, when, in particular, time plays a key role.
Animals↗
Abrupt learning and retinal size specificity in illusory-contour perception.
BACKGROUND: In behavioral studies of learning, a distinction is commonly made between gradual and abrupt improvements in performance. The learning of perceptual and motor skills is often characterized by gradual, incremental improvement, and is found not to generalize over stimulus manipulations such as changes in the size or location of the retinal image. In contrast, marked improvement in performance can occur suddenly - a phenomenon which has been termed 'insight'. Consequently, the brain mechanisms subserving the two types of learning are commonly thought of as distinct. Here, we examine learning of a perceptual task in which improvement appears to exhibit characteristics of both gradual and abrupt learning. RESULTS: We describe experiments on illusory-contour perception in which the observers underwent an abrupt, dramatic improvement in performance, resembling an incident of insight. At the same time, however, the phenomenon showed a degree of stimulus-specificity that was previously thought to characterize incremental, gradual learning. The improvement was triggered only by specific visual stimuli, whereas other, quite similar, stimuli were found to be ineffective for training; the learning did not generalize to a new retinal image size, and re-training was necessary for different-sized images. CONCLUSIONS: The juxtaposition of abrupt and stimulus-specific learning that we observed suggests that the distinction between the two forms of learning needs to be revised. Rather than postulating two distinct mechanisms, incremental and insightful learning need to be addressed within a single framework. In particular, the findings suggest that learning may involve interactions between multiple levels of representations of the stimulus.
Adolescent↗
Maladaptive anticipatory saccades in schizophrenia.
We compared the saccades made by 8 neuroleptic-treated and 7 drug-free schizophrenic inpatients with those made by 11 normal controls during two eye movement tasks. The first task was designed to elicit visually guided but not internally guided saccades. The second task was designed so that optimal performance required saccades be guided on the basis of an internal representation of target behavior. During the first task, schizophrenics made visually guided saccades that were as accurate as those made by control, but both drug-free and neuroleptic-treated schizophrenics made intrusive saccades at a significantly higher rate than control subjects. Most of these maladaptive saccades appeared to be premature attempts to anticipate target jump. During the second eye movement task, which for optimal performance required use of an internal representation to guide eye movements, most patients learned to anticipate target jump as well as controls. However, neuroleptic-treated patients made significantly smaller adaptive anticipatory saccades than either drug-free schizophrenic patients or normal subjects. These finding are discussed as they relate to the prefrontal cortex-basal ganglia circuits involved in the regulation of behavior by representational knowledge and the idea that the abnormal anticipatory saccades we observed represent a failure in the sensorimotor gating of information derived from internal representations.
Adult↗
Consolidation of visual associative long-term memory in the temporal cortex of primates.
Neuropsychological theories have proposed a critical role for the interaction between the medial temporal lobe and the neocortex in the formation of long-term memory for facts and events, which has often been tested by learning of a series of paired words or figures in humans. We have examined neural mechanisms underlying the memory "consolidation" process by single-unit recording and molecular biological methods in an animal model of a visual pair-association task in monkeys. In our previous studies, we found that long-term associative representations of visual objects are acquired through learning in the neural network of the anterior inferior temporal (IT) cortex. In this article, we propose the hypothesis that limbic neurons undergo rapid modification of synaptic connectivity and provide backward signals that guide the reorganization of neocortical neural circuits. Two experiments tested this hypothesis: (1) we examined the role of the backward connections from the medial temporal lobe to the IT cortex by injecting ibotenic acid into the entorhinal and perirhinal cortices, which provided massive backward projections ipsilaterally to the IT cortex. We found that the limbic lesion disrupted the associative code of the IT neurons between the paired associates, without impairing the visual response to each stimulus. (2) We then tested the first half of this hypothesis by detecting the expression of immediate-early genes in the monkey temporal cortex. We found specific expression of zif268 during the learning of a new set of paired associates in the pair-association task, most intensively in area 36 of the perirhinal cortex. All these results with the visual pair-association task support our hypothesis and demonstrate that the consolidation process, which was first proposed on the basis of clinico-psychological evidence, can now be examined in primates using neurophysiolocical and molecular biological approaches.
Analysis of Variance↗
Crossed unilateral lesions of the medial forebrain bundle and either inferior temporal or frontal cortex impair object-reward association learning in Rhesus monkeys.
In an accompanying paper we showed that combined transection of the fornix, amygdala and temporal stem in monkeys produced dense amnesia, including an impairment in visual object-reward association learning. We proposed that this combined surgical section had its effect by isolating temporal cortex from the ascending projections of the basal forebrain and midbrain structures. To test this hypothesis, in the present experiment we disconnected the inferior temporal cortex from these basal forebrain and midbrain structures, while sparing cortical white matter, by crossed unilateral lesions of the medial forebrain bundle in one hemisphere and inferior temporal cortex in the opposite hemisphere. The aim of the medial forebrain bundle lesion was to section axons of cells, both those that project to the cortex via the medial forebrain bundle, and those which control the activity of these same structures. A single unilateral lesion alone had no effect on the ability to learn and remember visual object-reward associations, but the crossed unilateral lesions produced an impairment in this task which was equal in severity to the impairment seen earlier after bilateral section of the fornix, amygdala and temporal stem. The impairment was not an effect of interrupting fibres to the cortex from the ventromedial hypothalamus, or of unilateral sensory neglect. This supports the hypothesis that these midbrain and basal forebrain afferents to the inferior temporal cortex are important for new visual learning. Furthermore, an impairment of equal severity was demonstrated in a separate group of animals that received crossed unilateral lesions of the medial forebrain bundle in one hemisphere and of the frontal cortex in the opposite hemisphere. We propose that the frontal cortex acts to modulate basal forebrain activity which in turn reinforces object representations in the inferior temporal cortex during learning.
Animals↗
Cortical remodelling induced by activity of ventral tegmental dopamine neurons.
Representations of sensory stimuli in the cerebral cortex can undergo progressive remodelling according to the behavioural importance of the stimuli. The cortex receives widespread projections from dopamine neurons in the ventral tegmental area (VTA), which are activated by new stimuli or unpredicted rewards, and are believed to provide a reinforcement signal for such learning-related cortical reorganization. In the primary auditory cortex (AI) dopamine release has been observed during auditory learning that remodels the sound-frequency representations. Furthermore, dopamine modulates long-term potentiation, a putative cellular mechanism underlying plasticity. Here we show that stimulating the VTA together with an auditory stimulus of a particular tone increases the cortical area and selectivity of the neural responses to that sound stimulus in AI. Conversely, the AI representations of nearby sound frequencies are selectively decreased. Strong, sharply tuned responses to the paired tones also emerge in a second cortical area, whereas the same stimuli evoke only poor or non-selective responses in this second cortical field in naive animals. In addition, we found that strong long-range coherence of neuronal discharge emerges between AI and this secondary auditory cortical area.
Acoustic Stimulation↗
The limitations of decision trees and automatic learning in real world medical decision making.
The decision tree approach is one of the most common approaches in automatic learning and decision making. It is popular for its simplicity in constructing, efficient use in decision making and for simple representation, which is easily understood by humans. The automatic learning of decision trees and their use usually show very good results in various "theoretical" environments. The training sets are usually large enough for learning algorithm to construct a hypothesis consistent with the underlying concept. But in real life it is often impossible to find the desired number of training objects for various reasons. The lack of possibilities to measure attribute values, high cost and complexity of such measurements, unavailability of all attributes at the same time are the typical representatives. There are different ways to deal with some of these problems, but in a delicate field of medical decision making, we cannot allow ourselves to make any inaccurate decisions. We have measured the values of 24 attributes before and after the 82 operations of children in age between 2 and 10 years. The aim was to find the dependencies between attribute values and a child's predisposition to acidemia--the decrease of blood's pH. Our main interest was in discovering predisposition to two forms of acidosis, the metabolic acidosis and the respiratory acidosis, which can both have serious effects on child's health. We decided to construct different decision trees from a set of training objects, which was complete (there were no missing attribute values), but on the other hand not large enough to avoid the effect of overfitting. A common approach to evaluation of a decision tree is the use of a test set. In our case we decided that instead of using a test set, we ask medical experts to take a closer look at the generated trees. They examined and evaluated the decision trees branch by branch. Their comments on the generated trees can be found in this paper. The comments show, that trees generated from available training set mainly have surprisingly good branches, but on the other hand some are very "stupid" and no medical explanation could be found. Thereafter we can conclude, that the decision tree concept and automatic learning can be successfully used in real world situations, constrained with the real world limitations, but they should be used only with the guidelines of appropriate medical experts.
Acidosis↗
Entorhinal cortex lesions disrupt the relational organization of memory in monkeys.
Recent accounts suggest that the hippocampal system critically supports two central characteristics of episodic memory: the ability to establish and maintain representations for the salient relationships between experienced events (relational representation) and the capacity to flexibly manipulate memory (flexible memory expression). To test this proposal in monkeys, intact controls and subjects with bilateral aspiration lesions of the entorhinal cortex were trained postoperatively on two standard memory tasks, delayed nonmatchingto-sample (DNMS) and two-choice object discrimination (OD) learning, and three procedures intended to emphasize relational representation and flexible memory expression: a paired associate (PA) task, a transitive inference (TI) test of learning and memory for hierarchical stimulus relationships, and a spatial delayed recognition span (SDRS) procedure. The latter assessments each included critical "probe" tests that asked monkeys to evaluate the relationships among previously learned stimuli presented in novel combinations. Subjects with entorhinal cortex lesions scored as accurately as controls on all phases of DNMS and OD, procedures that can be solved on the basis of memory for individual stimuli. In contrast, experimental monkeys displayed deficits relative to controls on all phases of the PA, TI, and SDRS tasks that emphasized the flexible manipulation of memory for the relationships between familiar items. Together, the findings support the conclusion that the primate hippocampal system critically enables the relational organization of declarative memory.
Animals↗
Neuronal tuning and associative mechanisms in form representation.
We examine the hypothesis that the form representation in the anterior inferotemporal (AIT) cortex is acquired through learning. According to this hypothesis, perceptual aspects of the temporal association area are closely related to its visual representation, in that the response selectivity of AIT neurons can be influenced by visual experience. On the basis of the neurophysiological evidence, we summarize two neuronal mechanisms that subserve the acquisition of form selectivity in AIT neurons. The first mechanism is neuronal tuning to particular stimuli that were learned in a cognitive task. The second mechanism is association, with which relevant information can be retrieved from other stored memories. On the grounds that long-term memory of objects is acquired and organized by at least these two neuronal mechanisms in the temporal association area, we further present a model of the cognitive memory system that unifies perception and imagery.
Animals↗
Speech sound representation in the brain.
Biologic processes underlying speech sound perception and learning have been addressed using the mismatch negativity (MMN) evoked response. First is a consideration of how the acoustic properties of the signal affect the neural mechanisms and brain regions engaged. Because the MMN differs depending on the acoustic characteristics of the stimuli used to elicit the response, it has been used to probe mechanisms underlying the neural representation of stimuli along the auditory pathway. Second is a consideration of neurophysiologic correlates of speech sound perception and learning. Detailed is a 'behavioral-neurophysiologic, acoustic-phonetic approach', used to link perception with underlying physiologic processes in humans. The focus here is on children and what has been learned about normal maturation of speech sound perception and its disruption in certain children with learning disorders. The last topic is a consideration of central nervous system changes with perceptual learning. This includes long-term experience with one's native language and short-term auditory training in the laboratory. Limitations and future challenges are discussed.
Auditory Pathways↗
Incidental concept learning, feature frequency, and correlated properties.
Four experiments examined sensitivity to feature frequencies and feature correlations as a function of intentional and incidental concept learning. Feature frequencies were encoded equally well across variations in learning strategies, and although classification decisions in both intentional and incidental conditions preserved correlated features, this sensitivity was achieved through different processes. With intentional learning, sensitivity to correlations resulted from explicit rules, whereas incidental encoding preserved correlations through a similarity-based analogical process. In incidental tasks that promoted exemplar storage, classification decisions were mediated by similarity to retrieved examples, and correlated features were indirectly preserved in this process. The results are discussed in terms of the diversity of encoding processes and representations that can occur with incidental category learning.
Adult↗
Activation of preexisting and acquired face representations: the N250 event-related potential as an index of face familiarity.
Electrophysiological studies using event-related potentials have demonstrated that face stimuli elicit a greater negative brain potential in right posterior recording sites 170 msec after stimulus onset (N170) relative to nonface stimuli. Results from repetition priming paradigms have shown that repeated exposures of familiar faces elicit a larger negative brainwave (N250r) at inferior temporal sites compared to repetitions of unfamiliar faces. However, less is known about the time course and learning conditions under which the N250 face representation is acquired. In the familiarization phase of the Joe/no Joe task, subjects studied a target "Joe" face ("Jane" for female subjects) and, during the course of the experiment, identified a series of sequentially presented faces as either Joe or not Joe. The critical stimulus conditions included the subject's own face, a same-sex Joe ( Jane) face and a same-sex "other" face. The main finding was that the subject's own face produced a focal negative deflection (N250) in posterior channels relative to nontarget faces. The task-relevant Joe target face was not differentiated from other nontarget faces in the first half of the experiment. However, in the second half, the Joe face produced an N250 response that was similar in magnitude to the own face. These findings suggest that the N250 indexes two types of face memories: a preexperimentally familiar face representation (i.e., the "own face" and a newly acquired face representation (i.e., the Joe/Jane face) that was formed during the course of the experiment.
Adolescent↗
Monkey perirhinal cortex is critical for visual memory, but not for visual perception: reexamination of the behavioural evidence from monkeys.
Overdependence on discrimination learning paradigms to assess the function of perirhinal cortex has complicated understanding of the cognitive role of this structure. Impairments in discrimination learning can result from at least two distinct causes: (a) failure to accurately apprehend and represent the relevant stimuli, or (b) failure to form and remember associations between stimulus representations and reward. Thus, the results of discrimination learning experiments do not readily differentiate deficits in perception from deficits in learning and memory. Here I describe studies that do dissociate learning and memory from perception and show that perirhinal cortex damage impairs learning and/or memory, but not perception. Reanalysis and reconsideration of other published data call into further question the hypothesis that the monkey perirhinal cortex plays a critical role in visual perception.
Animals↗
Inactivation of hippocampus or caudate nucleus with lidocaine differentially affects expression of place and response learning.
Involvement of the hippocampus and caudate nucleus in place and response learning was examined by functionally inactivating these brain regions bilaterally with infusions of lidocaine. Rats were trained to approach a consistently baited arm in a cross-maze from the same start box (four trials/day/14 total days). On Days 8 and 16 a single probe trial was given, in which rats were placed in the start box opposite that used in training and allowed to approach a maze arm. Three minutes prior to the probe trial, rats received bilateral injections of either saline or a 2% lidocaine solution (in order to produce neural inactivation) into either the dorsal hippocampus or dorsolateral caudate nucleus. On the probe trials, rats which entered the baited maze arm (i.e., approached the place where food was located during training) were designated place learners, and rats which entered the unbaited maze arm (i.e., made the same turning response as during training) were designated response learners. Saline-treated rats displayed place learning on the Day 8 probe trial and response learning on the Day 16 probe trial, indicating that with extended training there is a shift in learning mechanisms controlling behavior. Rats given lidocaine injections into the hippocampus showed no preference for place or response learning on the Day 8 probe trial, but displayed response learning on the Day 16 probe trial, indicating a blockade of place learning following inactivation of the hippocampus. Rats given lidocaine injections into the caudate nucleus displayed place learning on both the Day 8 and the Day 16 probe trials, indicating a blockade of response learning following inactivation of the caudate nucleus. The findings indicate: (1) the hippocampus and caudate nucleus selectively mediate expression of place and response learning, respectively (2), in a visually cued extramaze environment, hippocampal-dependent place learning is acquired faster than caudate-dependent response learning, and (3) when animals shift to caudate-dependent response learning with extended training, the hippocampal-based place representation remains intact.
Anesthetics↗
An analysis of the neural representation of birdsong memory.
Songbirds, such as zebra finches, learn their song from a tutor early in life. Forebrain nuclei in the "song system" are important for the acquisition and production of song. Brain regions [including the caudomedial part of the neostriatum (NCM) and of the hyperstriatum ventrale (CMHV)] outside the song system show increased neuronal activation, measured as expression of immediate early genes (IEGs), when zebra finch males are exposed to song. IEG expression in the NCM in response to tutor song is significantly positively correlated with the strength of song learning (i.e., the number of elements copied). Here, we exposed three groups of adult zebra finch males to tutor song, to their own song, or to novel conspecific song. The two control groups were included to examine an alternative explanation of our previous results in terms of variation in predisposed levels of attentiveness. Expression of Zenk, the protein product of the IEG ZENK, was measured in the NCM, CMHV, and hippocampus. There were no significant differences in overall Zenk expression between the three experimental groups. However, there was a significant positive correlation between Zenk expression in the NCM (but not in the other two regions) and strength of song learning in the males that were exposed to the tutor song. There was no such correlation in the other two groups. These results suggest that experience-related neuronal activation is specific to the tutor song and thus unlikely to be a result of differences in attention.
Analysis of Variance↗