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Representations of motivational drives in mesial cortex, medial thalamus, hypothalamus and midbrain.

We propose that neural representations of motivational drives, including sexual desire, hunger, thirst, fear, power-dominance, the motivational aspect of pain, the need for sleep, and nurturance, are represented in four areas in the brain. These are located in the medial hypothalamic/preoptic area, the periaqueductal gray matter (PAG) in the midbrain/pons, the midline and intralaminar thalamic nuclei, and in the anterior part of the mesial cortex, including the medial prefrontal and anterior cingulate areas. We attempt to determine the locations of each of these representations within the hypothalamus/preoptic area, periaqueductal gray and cortex, based on the available literature on activation of brain structures by stimuli that evoke these forms of motivation, on the effects of electrical and chemical stimulation and lesions of candidate structures, and on hodological data. We discuss the hierarchical organization of the representations for a given drive, outputs from these representations to premotor structures in the medulla, caudate-putamen, and cortex, and their contributions to involuntary, learned-sequential (operant) and voluntary behaviors.

Animals↗

Event-related potential indices of semantic priming following an unrelated intervening item.

Interposing an unrelated word between related primes and targets often disrupts priming. This finding has been used to support the view that semantic information is represented in a distributed fashion, rather than locally. In some studies where unrelated items intervened between the prime and target, however, significant priming was nevertheless obtained. The discrepant results of these studies has been attributed to differences in speed-accuracy tradeoff, post-lexical checking, conscious rehearsal of the prime and differences in the depth to which the prime and target were processed. The present study was designed in such a way as to minimize variability associated with post-lexical influences. The N400 component of the human event-related potential was used as a physiological index of the extent to which priming occurred with and without the interposition of an unrelated item. Priming effects on both the amplitude and latency of the N400 were rendered non-significant by the presence of an intervening unrelated word. The results are interpreted as tentative evidence that semantic representations are distributed.

Adolescent↗

Does size really matter--using a decision tree approach for comparison of three different databases from the medical field of acute appendicitis.

Decision trees have been successfully used for years in many medical decision making applications. Transparent representation of acquired knowledge and fast algorithms made decision trees one of the most often used symbolic machine learning approaches. This paper concentrates on the problem of separating acute appendicitis, which is a special problem of acute abdominal pain, from other diseases that cause acute abdominal pain by use of an decision tree approach. Early and accurate diagnosing of acute appendicitis is still a difficult and challenging problem in everyday clinical routine. An important factor in the error rate is poor discrimination between acute appendicitis and other diseases that cause acute abdominal pain. This error rate is still high, despite considerable improvements in history-taking and clinical examination, computer-aided decision-support, and special investigation such as ultrasound. We investigated three databases of different size with cases of acute abdominal pain to complete this task as successful as possible. The results show that the size of the database does not necessary directly influence the success of the decision tree built on it. Surprisingly we got the best results from the decision trees built on the smallest and the biggest database, where the database with medium size (relative to the other two) was not so successful. Despite this we were able to produce decision tree classifiers that were capable of producing correct decisions on test data sets with accuracy up to 84%, sensitivity to acute appendicitis up to 90%, and specificity up to 80% on the same test set.

Acute Disease↗

The universal SNARC effect: the association between number magnitude and space is amodal.

It is thought that number magnitude is represented in an abstract and amodal way on a left-to-right oriented mental number line. Major evidence for this idea has been provided by the SNARC effect (Dehaene, Bossini, & Giraux, 1993): responses to relatively larger numbers are faster for the right hand, those to smaller numbers for the left hand, even when number magnitude is irrelevant. The SNARC effect has been used to index automatic access to a central semantic and amodal magnitude representation. However, this assumption of modality independence has never been tested and it remains uncertain if the SNARC effect exists in other modalities in a similar way as in the visual modality. We have examined this question by systematically varying modality/notation (auditory number word, visual Arabic numeral, visual number word, visual dice pattern) in a within-participant design. The SNARC effect was found consistently for all modality/notation conditions, including auditory presentation. The size of the SNARC effect in the auditory condition did not differ from the SNARC effect in any visual condition. We conclude that the SNARC effect is indeed a general index of a central semantic and amodal number magnitude representation.

Adolescent↗

Detection of three-dimensional structure in moving optical patterns.

Three experiments examined parameters affecting human observers' ability to detect the global three-dimensional (3D) organization of a random-dot display corresponding to the polar projection of a rotating sphere. Results indicate that the detection of 3D structure is critically dependent on the detectability of motion, is disrupted by increased redundant information specifying the two-dimensional location of points in the display, and undergoes a rapidly increasing resistance to the disruptive effects of noise with increasing numbers of frames. These results, in conjunction with earlier findings, are inconsistent with existing theories concerning the perception of three-dimensionality in moving displays, in that they indicate a high degree of visual sensitivity to stimulus organizations with unique topological representations.

Attention↗

Generation of visual representations.

The visual confusability of uppercase letters was manipulated in a successive same-different task to study the conditions under which visual generation from auditory inputs would occur and to investigate the figural specificity of the generated representations. Prior experiments have shown that visual confusions do occur when the initial stimulus is auditory and the second one is visual, which indicates that auditory stimuli can be encoded into visual forms. There has been some suggestion, however, that the generated visual code may have been too abstract to differentiate between the two cases in which letters can appear. In the present experiment, although the confusion effect was not eliminated when the subjects had no advance knowledge regarding the case in which the visual stimulus would appear, the marked confusion effect obtained when the visual stimulus was an uppercase letter was substantially attenuated when the letter appeared in lowercase. This was taken to indicate that the visual characteristics of a generated visual representation may be relatively specific. The results also suggested that subjects may wait until after the second stimulus is presented before they generate the visual representation of the initial auditory stimulus.

Discrimination Learning↗

Neural codes: firing rates and beyond.

Computational neuroscience has contributed significantly to our understanding of higher brain function by combining experimental neurobiology, psychophysics, modeling, and mathematical analysis. This article reviews recent advances in a key area: neural coding and information processing. It is shown that synapses are capable of supporting computations based on highly structured temporal codes. Such codes could provide a substrate for unambiguous representations of complex stimuli and be used to solve difficult cognitive tasks, such as the binding problem. Unsupervised learning rules could generate the circuitry required for precise temporal codes. Together, these results indicate that neural systems perform a rich repertoire of computations based on action potential timing.

Action Potentials↗

Contextual modulation of orientation tuning contributes to efficient processing of natural stimuli.

It has been proposed that sensory neurons are adapted to the statistical structure of the natural environment in order to encode natural stimuli efficiently. While spatiotemporal correlations in luminance signals may be decorrelated by neurons in early visual processing stages, higher-order correlations, such as those in the orientation domain, are likely to persist in the input representation until the cortical level. In this study, we first examine orientation correlations in natural stimuli across brief time intervals and across nearby regions of space, and find strong correlations in both domains. We then examine contextual modulation of orientation tuning. We find that both temporal and spatial contexts exert a common influence on orientation tuning, shifting tuning away from the orientation of either the adapting (temporal) or surrounding (spatial) grating. Finally, we incorporate this context-mediated repulsive shift in orientation tuning into a model of cortical responses. We find that a direct result of the shift is a reduction of the redundancy in the population responses evoked by the orientation configurations that are most common in natural stimuli. Thus, cortical neurons may be adapted to the statistics of orientation in natural stimuli in order to increase the efficiency of natural stimulus representation.

Animals↗

What differentiates declarative and procedural memories: reply to Cohen, Poldrack, and Eichenbaum (1997)

CPE claim that procedural and declarative representations differ on two important dimensions: flexibility and compositionality. I have proposed that the apparent flexibility of a memory depends entirely on the transfer conditions. Any retest is, in some sense, a test of flexibility, because something has changed since the original encoding episodic. I have argued that if one changes something that does not provide support to memory performance, the memory will appear flexible, and resistant to changes in the environment. If one changes the very thing that the representation codes, the memory will appear inflexible and easily disrupted by changes in the environment. This principle is equally true for procedural and declarative memory. CPE contend that procedural representations lack compositionality. An ideal test of this claim would examine the representation of a task that is widely agreed to be procedural (e.g. that has been demonstrated to be learned normally by amnesic patients, and in the absence of awareness by neurologically intact subjects). Such experiments appear not to have been conducted, and the fact is that many tasks that are widely agreed to be procedural probably are not compositional. They appear to be, as CPE contend, biases in a processing system; it is hard to imagine how repetition priming could be compositional. Nevertheless, this is not true of all procedural memories. There is a good deal of evidence that motor behaviour is organised hierarchically and has compositionality. There is every reason to think that most if not all motor behaviour is procedural; motor behaviour might be driven by goals that are declarative, but the low-level operations that actually manipulate effectors are closed to consciousness, do not depend on the medial temporal lobe or diencephalon, and would therefore be classified as procedural. CPE framed their theory of differences between procedural and declarative memory systems as an account of the deficit in amnesic patients. They therefore predict that the learning of amnesic patients should not show flexibility or compositionality. There is already at least one study showing learning in amnesic patients that is as flexible as that of control participants (Knowlton & Squire, 1996). There are not, to my knowledge, data on whether the motor skill learning of amnesic patients shows compositionality, but one might expect that it would, given that it does in neurologically intact participants, and given that motor skill learning appears unimpaired in amnesic patients. Thus, the conception of declarative and procedural memory provided by CPE may not provide a complete account of amnesic performance. The anatomic distinction between procedural and declarative memory systems appears quite strong, and there is therefore reason to believe that there are accompanying computational differences. There does not, however, appear to be sufficient evidence to support those differences proposed by CPE.

Amnesia↗

A global bioheat model with self-tuning optimal regulation of body temperature using Hebbian feedback covariance learning.

In the lower brain, body temperature is continually being regulated almost flawlessly despite huge fluctuations in ambient and physiological conditions that constantly threaten the well-being of the body. The underlying control problem defining thermal homeostasis is one of great enormity: Many systems and sub-systems are involved in temperature regulation and physiological processes are intrinsically complex and intertwined. Thus the defining control system has to take into account the complications of nonlinearities, system uncertainties, delayed feedback loops as well as internal and external disturbances. In this paper, we propose a self-tuning adaptive thermal controller based upon Hebbian feedback covariance learning where the system is to be regulated continually to best suit its environment. This hypothesis is supported in part by postulations of the presence of adaptive optimization behavior in biological systems of certain organisms which face limited resources vital for survival. We demonstrate the use of Hebbian feedback covariance learning as a possible self-adaptive controller in body temperature regulation. The model postulates an important role of Hebbian covariance adaptation as a means of reinforcement learning in the thermal controller. The passive system is based on a simplified 2-node core and shell representation of the body, where global responses are captured. Model predictions are consistent with observed thermoregulatory responses to conditions of exercise and rest, and heat and cold stress. An important implication of the model is that optimal physiological behaviors arising from self-tuning adaptive regulation in the thermal controller may be responsible for the departure from homeostasis in abnormal states, e.g., fever. This was previously unexplained using the conventional "set-point" control theory.

Adaptation, Physiological↗

Computational studies on acquisition and adaptation of ocular following responses based on cerebellar synaptic plasticity.

To investigate how cerebellar synaptic plasticity guides the acquisition and adaptation of ocular following response (OFR), a large-scale network model was developed. The model includes the cerebral medial superior temporal area (MST), Purkinje cells (P cells) of the ventral paraflocculus, the accessory optic and climbing fiber systems, the brain stem oculomotor network, and the oculomotor plant. The model reconstructed temporal profiles of both firing patterns of MST neurons and P cells and eye movements. Model MST neurons (n = 1,080) were set to be driven by retinal error and exhibited 12 preferred directions, 30 preferred velocities, and 3 firing waveforms. Correspondingly, each model P cell contained 1,080 excitatory synapses from granule cell axons (GCA) and 1,080 inhibitory synapses. P cells (n = 40) were classified into four groups by their laterality (hemisphere) and by preferred directions of their climbing fiber inputs (CF) (contralateral or upward). The brain stem neural circuit and the oculomotor plant were modeled on the work of Yamamoto et al. The initial synaptic weights on the P cells were set randomly. At the beginning, P cell simple spikes were not well modulated by visual motion, and the eye was moved only slightly by the accessory optic system. The synaptic weights were updated according to integral-differential equation models of physiologically demonstrated synaptic plasticity: long-term depression and long-term potentiation for GCA synapses and rebound potentiation for inhibitory synapses. We assumed that maximum plasticity was induced when GCA inputs preceded CF inputs by 200 ms. After more than 10,000 presentations of ramp-step visual motion, the strengths of both the excitatory and inhibitory synapses were modified. Subsequently, the simple spike responses became well developed, and ordinary OFRs were acquired. The preferred directions of simple spikes became the opposite of those of CFs. Although the model MST neurons were set to possess a wide variety of firing characteristics, the model P cells acquired only downward or ipsilateral preferred directions, high preferred velocities and stereotypical firing waveforms. Therefore the drastic transition of the neural representation from the population codes in the MST to the firing-rate codes of simple spikes were learned at the GCA-P cell synapses and inhibitory cells-P cell synapses. Furthermore, the model successfully reproduced the gain- and directional-adaptation of OFR, which was demonstrated by manipulating the velocity and direction of visual motion, respectively. When we assumed that synaptic plasticity could only occur if CF inputs preceded GCA inputs, the ordinary OFR were acquired but neither the gain-adaptation nor the directional adaptation could be reproduced.

Adaptation, Physiological↗

Remapping hand movements in a novel geometrical environment.

The issue of how the Euclidean properties of space are represented in the nervous system is a main focus in the study of visual perception, but is equally relevant to motor learning. The goal of our experiments was to investigate how the properties of space guide the remapping of motor coordination. Subjects wore an instrumented data glove that recorded the finger motions. Signals generated by the glove operated a remotely controlled endpoint: a cursor on a computer monitor. The subjects were instructed to execute movements of this endpoint with controlled motions of the fingers. This required inverting a highly redundant map from fingers to cursor motions. We found that 1) after training with visual feedback of the final error (but not of the ongoing cursor motion), subjects learned to map cursor locations into configurations of the fingers; 2) extended practice of movement led to more rectilinear cursor movement, a trend facilitated by training under continuous visual feedback of cursor motions; 3) with practice, subjects reduced motion in the degrees of freedom that did not contribute to the movements of the cursor; 4) with practice, subjects reduced variability of both cursor and hand movements; and 5) the reduction of errors and the increase in linearity generalized beyond the set of movements used for training. These findings suggest that subjects not only learned to produce novel coordinated movement to control the placement of the cursor, but they also developed a representation of the Euclidean space on which hand movements were remapped.

Analysis of Variance↗

Structure and function of learning flights in ground-nesting bees and wasps

Bees and wasps perform systematic flight manoevres when they leave their nest or a foodplace, during which they acquire or update their visual memory of the goal location. In a typical learning flight, the insect backs away from the goal in a series of arcs that are roughly centred on the goal. The mean rate of turning is rather constant and tends to balance the angular speed at which the arc is described. As a result, the insect views the goal at relatively fixed retinal positions in its left and right visual field, depending on flight direction. The general direction in which the insect backs away from the goal and the transition from one arc segment to the next are influenced by the local scene and by compass cues. Insects returning to the goal repeat some of the flight manoeuvres of their preceding learning flights. Their orientation in space and the retinal positions at which they view nearby landmarks are similar. One important function of learning flights appears to be the acquisition of visual depth information. We review the consequences of the structure of learning flights for visual information processing and discuss how they may relate to the acquisition of a visual representation and the task of pinpointing the goal.

Journal Article↗

Adaptivity of tuning functions in a generic recurrent network model of a cortical hypercolumn.

The representation of orientation information in the adult visual cortex is plastic as exemplified by phenomena such as perceptual learning or attention. Although these phenomena operate on different time scales and give rise to different changes in the response properties of neurons, both lead to an improvement in visual discrimination or detection tasks. If, however, optimal performance is indeed the goal, the question arises as to why the changes in neuronal response properties are so different. Here, we hypothesize that these differences arise naturally if optimal performance is achieved by means of different mechanisms. To evaluate this hypothesis, we set up a recurrent network model of a visual cortical hypercolumn and asked how each of four different parameter sets (strength of afferent and recurrent synapses, neuronal gains, and additive background inputs) must be changed to optimally improve the encoding accuracy of a particular set of visual stimuli. We find that the predicted changes in the population responses and the tuning functions were different for each set of parameters, hence were strongly dependent on the plasticity mechanism that was operative. An optimal change in the strength of the recurrent connections, for example, led to changes in the response properties that are similar to the changes observed in perceptual learning experiments. An optimal change in the neuronal gains led to changes mimicking neural effects of attention. Assuming the validity of the optimal encoding hypothesis, these model predictions can be used to disentangle the mechanisms of perceptual learning, attention, and other adaptation phenomena.

Animals↗

Informational primacy of visual dimensions: specialized roles for luminance and chromaticity in figure-ground perception.

Three experiments were conducted to examine the distinct contributions of two visual dimensions to figure-ground segregation. In each experiment, pattern identification was assessed by asking observers to judge whether a near-threshold test pattern was the same or different in shape to a high-contrast comparison pattern. A test pattern could differ from its background along one dimension, either luminance (luminance tasks) or chromaticity (chromaticity tasks). In each task, performance in a baseline condition, in which the test pattern was intact, was compared with performance in each of several degradation conditions, in which either the contour or the surface of the figure was degraded, using either partial occlusion (Experiment 1) or ramping (Experiments 2 and 3) of figure-ground differences. In each experiment, performance in luminance tasks was worst when the contour was degraded, whereas performance in chromaticity tasks was worst when the surface was degraded. This interaction was found even when spatial frequencies were fixed across test patterns by low-pass filtering. The results are consistent with a late (postfiltering) dual-mechanism system that processes luminance information to extract boundary representations and chromaticity information to extract surface representations.

Attention↗

Categorizing chairs and naming pears: category differences in object processing as a function of task and priming.

Four experiments are reported examining the locus of structural similarity effects in picture recognition and naming with normal subjects. Subjects carried out superordinate categorization and naming tasks with picture and word forms of clothing, furniture, fruit, and vegetable exemplars. The main findings were as follows: (1) Responses to pictures of fruit and vegetables ("structurally similar" objects) were slowed relative to pictures of clothing and furniture ("structurally dissimilar" objects). This structural similarity difference was greater for picture naming than for superordinate categorization of pictures. (2) Structural similarity effects in picture naming were reduced by repetition priming. Repetition priming effects were equivalent from picture and word naming as prime tasks. (3) However, superordinate categorization of the prime did not produce the structural similarity effects on priming found for picture naming. Furthermore, such priming effects did not arise for picture or word categorization or for reading picture names as target tasks. It is proposed that structural similarity effects on priming object processing are located in processes mapping semantic representations of pictures to name representations required to select names for objects. Visually based competition between fruit and vegetables produces competition in name selection, which is reduced by priming the mappings between semantic and name representations.

Adult↗

Memory for objects and parts.

Performance on whole comparisons of two whole objects was compared to that for partial comparisons of a whole object and a part. In particular, same-different comparisons were measured for sequential displays of two-dimensional line figures. The comparisons were analogous to those in the word priority paradigm (Johnson, 1975). The whole comparisons were found to be more accurate and faster than the partial comparisons. This whole advantage was also found for accuracy using a two-alternative forced-choice procedure. This effect was found for brief displays of a single object and for longer duration displays of two objects. It was also found for three different sets of line figures. In contrast, the whole advantage was not found for unconnected figures. These findings were interpreted in the context of hierarchical representations of objects and parts.

Adult↗