Biomedical subjects

E T Rolls

Publications and source records attributed to E T Rolls.

At least 55 records · Page 3Linked to original sources

PubMed2000

Memory systems in the brain.

The operation of different brain systems involved in different types of memory is described. One is a system in the primate orbitofrontal cortex and amygdala involved in representing rewards and punishers, and in learning stimulus-reinforcer associations. This system is involved in emotion and motivation. A second system in the temporal cortical visual areas is involved in learning invariant representations of objects. A third system in the hippocampus is implicated in episodic memory and in spatial function. Fourth, brain systems in the frontal and temporal cortices involved in short term memory are described. The approach taken provides insight into the neuronal operations that take place in each of these brain systems, and has the aim of leading to quantitative biologically plausible neuronal network models of how each of these memory systems actually operates.

Animals↗

PubMed2000

A model of invariant object recognition in the visual system: learning rules, activation functions, lateral inhibition, and information-based performance measures.

VisNet2 is a model to investigate some aspects of invariant visual object recognition in the primate visual system. It is a four-layer feedforward network with convergence to each part of a layer from a small region of the preceding layer, with competition between the neurons within a layer and with a trace learning rule to help it learn transform invariance. The trace rule is a modified Hebbian rule, which modifies synaptic weights according to both the current firing rates and the firing rates to recently seen stimuli. This enables neurons to learn to respond similarly to the gradually transforming inputs it receives, which over the short term are likely to be about the same object, given the statistics of normal visual inputs. First, we introduce for VisNet2 both single-neuron and multiple-neuron information-theoretic measures of its ability to respond to transformed stimuli. Second, using these measures, we show that quantitatively resetting the trace between stimuli is not necessary for good performance. Third, it is shown that the sigmoid activation functions used in VisNet2, which allow the sparseness of the representation to be controlled, allow good performance when using sparse distributed representations. Fourth, it is shown that VisNet2 operates well with medium-range lateral inhibition with a radius in the same order of size as the region of the preceding layer from which neurons receive inputs. Fifth, in an investigation of different learning rules for learning transform invariance, it is shown that VisNet2 operates better with a trace rule that incorporates in the trace only activity from the preceding presentations of a given stimulus, with no contribution to the trace from the current presentation, and that this is related to temporal difference learning.

Algorithms↗

PubMed1999

On decoding the responses of a population of neurons from short time windows.

The effectiveness of various stimulus identification (decoding) procedures for extracting the information carried by the responses of a population of neurons to a set of repeatedly presented stimuli is studied analytically, in the limit of short time windows. It is shown that in this limit, the entire information content of the responses can sometimes be decoded, and when this is not the case, the lost information is quantified. In particular, the mutual information extracted by taking into account only the most likely stimulus in each trial turns out to be, if not equal, much closer to the true value than that calculated from all the probabilities that each of the possible stimuli in the set was the actual one. The relation between the mutual information extracted by decoding and the percentage of correct stimulus decodings is also derived analytically in the same limit, showing that the metric content index can be estimated reliably from a few cells recorded from brief periods. Computer simulations as well as the activity of real neurons recorded in the primate hippocampus serve to confirm these results and illustrate the utility and limitations of the approach.

Algorithms↗

PubMed1999

Backward projections in the cerebral cortex: implications for memory storage.

Cortical areas are characterized by forward and backward connections between adjacent cortical areas in a processing stream. Within each area there are recurrent collateral connections between the pyramidal cells. We analyze the properties of this architecture for memory storage and processing. Hebb-like synaptic modifiability in the connections and attractor states are incorporated. We show the following: (1) The number of memories that can be stored in the connected modules is of the same order of magnitude as the number that can be stored in any one module using the recurrent collateral connections, and is proportional to the number of effective connections per neuron. (2) Cooperation between modules leads to a small increase in memory capacity. (3) Cooperation can also help retrieval in a module that is cued with a noisy or incomplete pattern. (4) If the connection strength between modules is strong, then global memory states that reflect the pairs of patterns on which the modules were trained together are found. (5) If the intermodule connection strengths are weaker, then separate, local memory states can exist in each module. (6) The boundaries between the global and local retrieval states, and the nonretrieval state, are delimited. All of these properties are analyzed quantitatively with the techniques of statistical physics.

Cerebral Cortex↗

PubMed1999

Correlations and the encoding of information in the nervous system.

Is the information transmitted by an ensemble of neurons determined solely by the number of spikes fired by each cell, or do correlations in the emission of action potentials also play a significant role? We derive a simple formula which enables this question to be answered rigorously for short time-scales. The formula quantifies the corrections to the instantaneous information rate which result from correlations in spike emission between pairs of neurons. The mutual information that the ensemble of neurons conveys about external stimuli can thus be broken down into firing rate and correlation components. This analysis provides fundamental constraints upon the nature of information coding, showing that over short time-scales correlations cannot dominate information representation, that stimulus-independent correlations may lead to synergy (where the neurons together convey more information than they would if they were considered independently), but that only certain combinations of the different sources of correlation result in significant synergy rather than in redundancy or in negligible effects. This analysis leads to a new quantification procedure which is directly applicable to simultaneous multiple neuron recordings.

Action Potentials↗

PubMed1999

Firing rate distributions and efficiency of information transmission of inferior temporal cortex neurons to natural visual stimuli.

The distribution of responses of sensory neurons to ecological stimulation has been proposed to be designed to maximize information transmission, which according to a simple model would imply an exponential distribution of spike counts in a given time window. We have used recordings from inferior temporal cortex neurons responding to quasi-natural visual stimulation (presented using a video of everyday lab scenes and a large number of static images of faces and natural scenes) to assess the validity of this exponential model and to develop an alternative simple model of spike count distributions. We find that the exponential model has to be rejected in 84% of cases (at the p < 0.01 level). A new model, which accounts for the firing rate distribution found in terms of slow and fast variability in the inputs that produce neuronal activation, is rejected statistically in only 16% of cases. Finally, we show that the neurons are moderately efficient at transmitting information but not optimally efficient.

Animals↗

PubMed1999

The representation of pleasant touch in the brain and its relationship with taste and olfactory areas.

Although there has been much investigation of brain pathways involved in pain, little is known about the brain mechanisms involved in processing somatosensory stimuli which feel pleasant. Employing fMRI it was shown that pleasant touch to the hand with velvet produced stronger activation of the orbitofrontal cortex than affectively neutral touch of the hand with wood. In contrast, the affectively neutral but more intense touch produced more activation of the primary somatosensory cortex than the pleasant stimulus. This indicates that part of the orbitofrontal cortex is concerned with representing the positively affective aspects of somatosensory stimuli, and in further experiments it was shown that this orbitofrontal area is different from that activated by taste and smell. The finding that three different primary or unlearned types of reinforcer (touch, taste, and smell) are represented in the orbitofrontal cortex helps to provide a firm foundation for understanding the neural basis of emotions, which can be understood in terms of states elicited by stimuli which are rewarding or punishing.

Brain↗

PubMed1999

Responses to the sensory properties of fat of neurons in the primate orbitofrontal cortex.

The primate orbitofrontal cortex is a site of convergence of information from primary taste, olfactory, and somatosensory cortical areas. We describe the responses of a population of single neurons in the orbitofrontal cortex that responds to fat in the mouth. The neurons respond, when fatty foods are being eaten, to pure fat such as glyceryl trioleate and also to substances with a similar texture but different chemical composition such as paraffin oil (hydrocarbon) and silicone oil [Si(CH3)2O)n]. This is evidence that the neurons respond to the oral texture of fat, sensed by the somatosensory system. Some of the population of neurons respond unimodally to the texture of fat. Other single neurons show convergence of taste inputs, and others of olfactory inputs, onto single neurons that respond to fat. For example, neurons were found that responded to the mouth feel of fat and the taste of monosodium glutamate (both found in milk), or to the mouth feel of fat and to odor. Feeding to satiety reduces the responses of these neurons to the fatty food eaten, but the neurons still respond to some other foods that have not been fed to satiety. Thus sensory-specific satiety for fat is represented in the responses of single neurons in the primate orbitofrontal cortex. Fat is an important constituent of food that affects its palatability and nutritional effects. The findings described provide evidence that the reward value (or pleasantness) of the mouth feel of fat is represented in the primate orbitofrontal cortex and that the representation is relevant to appetite.

Animal Nutritional Physiological Phenomena↗

PubMed1999

Head direction cells in the primate pre-subiculum.

The function of the primate hippocampus and related structures was analysed by making recordings from the hippocampus, subiculum, presubiculum, and parahippocampal gyrus in monkeys actively walking in the laboratory. Head direction cells were found in the presubiculum. The firing rate of these cells was a function of the head direction of the monkey, with a response that was typically 10-100 times larger to the best as compared to the opposite direction. The mean half-amplitude width of the tuning of the cells was 76 degrees. The response of head direction cells in the presubiculum was not influenced by the place where the monkey was, there being the same tuning to head direction at different places in a room, and even outside the room. The response of these cells was also independent of the "spatial view" observed by the monkey, and also the position of the eyes in the head. The average information about head direction was 0.64 bits, about place was 0.10 bits, about spatial view was 0.27 bits, and about eye position was 0.04 bits. The cells maintained their tuning for periods of at least several minutes when the view details were obscured or the room was darkened. This representation of head direction could be useful together with the hippocampal spatial view cells and whole body motion cells found in primates in such spatial and memory functions as path integration.

Action Potentials↗

PubMed1999

Spatial view cells and the representation of place in the primate hippocampus.

The information represented in the primate hippocampus is being analysed by making recordings in monkeys actively walking in the laboratory. In a sample of 352 cells recorded in this situation, no "place" cells have so far been found. Instead, we have found a considerable population of "spatial view" cells tuned to respond when the monkey looks at small parts of the environment. We have been able to demonstrate (1) that these hippocampal neurons respond to a view of space "out there," not to the place where the monkey is; (2) that the responses depend on where the monkey is looking, by measuring eye position; (3) that the responses in some cases (e.g., CA1 but not CA3) still occur if the view details are obscured with curtains; (4) that the cells (in, e.g., CA1) retain part of their "space" tuning even in complete darkness, for several minutes; and (5) that the spatial representation is allocentric. The spatial representation is, thus, different from that in the rat hippocampus, in which place cells respond based on where the rat is located. The representation is also different from that described in the parietal cortex, where neurons respond in egocentric coordinates. This representation of space "out there" provided by primate spatial view cells would be an appropriate part of a memory system involved in memories of particular events or episodes, for example, of where in an environment an object was seen. Spatial view cells (in conjunction with whole body motion cells in the primate hippocampus, and head direction cells in the primate presubiculum) would also be useful as part of a spatial navigation system, for which they would provide a memory component.

Animals↗

PubMed1999

Analysis of information transmission in the Schaffer collaterals.

Hippocampal region CA1 seems from comparative studies to be particularly important in the primate brain, in addition to being crucial to memory function. Thus, it is an extremely appropriate place to begin a quantitative investigation of the information representation and transmission capabilities of cerebral neural networks. In this study, a mathematical model of the Schaffer collateral projection from CA3 to CA1 is described. From the model, the amount of information that can be conveyed by the Schaffer collaterals is calculated, i.e., the information that a pattern of firing in CA1 conveys about a pattern of firing in CA3, because of the connections between them. The calculation is performed analytically for an arbitrary probability distribution describing the pattern of CA3 firing and then solved numerically for particular input distributions. The effect of a number of issues on the information conveyed is examined. Consideration of the effect of the amount of analog resolution of firing rates in the patterns of activity in CA3 confirmed information transmission to be most efficient for binary codes, to a degree that depends on the sparseness of activity. For very sparse codes, a binary code allows more information to be received even in absolute terms, but for more distributed codes, slightly more information can be received by CA1 by making use of analog resolution. The pattern of convergence of connections from CA3 to CA1 is examined, i.e., the spatial distribution of the number of connections each CA1 neuron receives. It is found that the effect of the difference between a uniform convergence model and a proposed real convergence pattern (Bernard and Wheal, Hippocampus 1994;4:497-529) is minimal. The effect of the ratio of expansion between CA3 and CA1 due to the relative numbers of neurons in these two areas is studied. The Schaffer collaterals in all mammalian species reported in the literature seem to operate in a regime in which there is at least the scope for efficient transfer of information. In addition, the effect of topography (with respect to the transverse hippocampal axis) in the Schaffer collateral connectivity is examined. In the absence of spatial correlations, topography is found to have essentially no effect on information transmission. If spatial correlations in firing were present in CA3 (which, however, would be less efficient for memory storage in the recurrent collaterals), information transmission would be maximized by matching the topographic spread to the spatial scale of correlation.

Animals↗

PubMed1999

The neurophysiology of backward visual masking: information analysis.

Backward masking can potentially provide evidence of the time needed for visual processing, a fundamental constraint that must be incorporated into computational models of vision. Although backward masking has been extensively used psychophysically, there is little direct evidence for the effects of visual masking on neuronal responses. To investigate the effects of a backward masking paradigm on the responses of neurons in the temporal visual cortex, we have shown that the response of the neurons is interrupted by the mask. Under conditions when humans can just identify the stimulus, with stimulus onset asynchronies (SOA) of 20 msec, neurons in macaques respond to their best stimulus for approximately 30 msec. We now quantify the information that is available from the responses of single neurons under backward masking conditions when two to six faces were shown. We show that the information available is greatly decreased as the mask is brought closer to the stimulus. The decrease is more marked than the decrease in firing rate because it is the selective part of the firing that is especially attenuated by the mask, not the spontaneous firing, and also because the neuronal response is more variable at short SOAs. However, even at the shortest SOA of 20 msec, the information available is on average 0.1 bits. This compares to 0.3 bits with only the 16-msec target stimulus shown and a typical value for such neurons of 0.4 to 0.5 bits with a 500-msec stimulus. The results thus show that considerable information is available from neuronal responses even under backward masking conditions that allow the neurons to have their main response in 30 msec. This provides evidence for how rapid the processing of visual information is in a cortical area and provides a fundamental constraint for understanding how cortical information processing operates.

Animals↗

PubMed1999

Associative memory properties of multiple cortical modules.

The existence of recurrent collateral connections between pyramidal cells within a cortical area and, in addition, reciprocal connections between connected cortical areas, is well established. In this work we analyse the properties of a tri-modular architecture of this type in which two input modules have convergent connections to a third module (which in the brain might be the next module in cortical processing or a bi-modal area receiving connections from two different processing pathways). Memory retrieval is analysed in this system which has Hebb-like synaptic modifiability in the connections and attractor states. Local activity features are stored in the intra-modular connections while the associations between corresponding features in different modules present during training are stored in the inter-modular connections. The response of the network when tested with corresponding and contradictory stimuli to the two input pathways is studied in detail. The model is solved quantitatively using techniques of statistical physics. In one type of test, a sequence of stimuli is applied, with a delay between them. It is found that if the coupling between the modules is low a regime exists in which they retain the capability to retrieve any of their stored features independently of the features being retrieved by the other modules. Although independent in this sense, the modules still influence each other in this regime through persistent modulatory currents which are strong enough to initiate recall in the whole network when only a single module is stimulated, and to raise the mean firing rates of the neurons in the attractors if the features in the different modules are corresponding. Some of these mechanisms might be useful for the description of many phenomena observed in single neuron activity recorded during short term memory tasks such as delayed match-to-sample. It is also shown that with contradictory stimulation of the two input modules the model accounts for many of the phenomena observed in the McGurk effect, in which contradictory auditory and visual inputs can lead to misperception.

Action Potentials↗

PubMed1998

The neurophysiology of taste and olfaction in primates, and umami flavor.

To investigate the neural encoding of glutamate (umami) taste in the primate, recordings were made from taste responsive neurons in the cortical taste areas in macaques. Most of the neurons were in the orbitofrontal cortex (secondary) taste area. First, it was shown that there is a representation of the taste of glutamate which is separate from the representation of the other prototypical tastants sweet (glucose), salt (NaCl), bitter (quinine) and sour (HCl). Second, it was shown that single neurons that had their best responses to sodium glutamate also had good responses to glutamic acid. Third, it was shown that the responses of these neurons to the nucleotide umami tastant inosine 5'-monophosphate were more correlated with their responses to monosodium glutamate than to any prototypical tastant. Fourth, concentration response curves showed that concentrations of monosodium glutamate as low as 0.001 M were just above threshold for some of these neurons. Fifth, some neurons in the orbitofrontal region, which responded to monosodium glutamate and other food tastes, decreased their responses after feeding with monosodium glutamate to behavioral satiety, revealing a mechanism of satiety. In some cases this reduction was sensory-specific. Sixth, it was shown in psychophysical experiments in humans that the flavor of umami is strongest with a combination of corresponding taste and olfactory stimuli (e.g., monosodium glutamate and garlic odor). The hypothesis is proposed that part of the way in which glutamate works as a flavor enhancer is by acting in combination with corresponding food odors. The appropriate associations between the odor and the glutamate taste may be learned at least in part by olfactory to taste association learning in the primate orbitofrontal cortex.

Animals↗

PubMed1998

View-invariant representations of familiar objects by neurons in the inferior temporal visual cortex.

A view-invariant representation of objects in the brain would have many computational advantages. Here we describe a population of single neurons in the temporal visual cortex (IT) that have view-invariant representations of familiar objects. Ten real plastic objects were placed in the monkeys' home cages for a period of time before neurophysiological experiments in which neuronal responses were measured to four views of each object. The macaques performed a visual fixation task, and had never been trained in object discrimination. The majority of the visual neurons recorded were responsive to some views of some objects and/or to the control stimuli, as would be expected from previous studies. However, a small subset of these neurons were responsive to all views of one or more of the objects, providing evidence that these neurons were coding for objects, rather than simply for individual views or visual features within the image. This result was confirmed by information theoretic analyses, which showed that the neurons provided information about which object was being seen, independently of the view. The coding scheme was shown to be sparse distributed, with relatively independent information being provided by the different neurons. Hypotheses about how these view-invariant cells are formed are described.

Action Potentials↗

PubMed1998

Spatial view cells in the primate hippocampus: effects of removal of view details.

Hippocampal function was analyzed by making recordings from hippocampal formation neurons in macaques actively walking in the laboratory. "Spatial view" cells, which respond when the monkey looks at a part of the environment were analyzed. It is shown that many of these cells retain their spatial characteristics when the view details are obscured totally by curtains and by darkness. It is shown that many of these cells respond more when the monkey is gazing toward one location in the room than toward other locations, even though none of the view details can be seen. Such cells were found in the CA1 region, the parahippocampal gyrus, and the presubiculum. Other cells stopped responding when the monkey looked toward the normally effective location in the environment if the view details were obscured. These cells were in the CA3 region of the hippocampus. The results indicate that for CA3 cells, the visual input is necessary for the normal spatial response of the neurons, and for other cells in the primate hippocampal formation, the response still depends on the monkey gazing toward that location in space when the view details are obscured. These latter cells therefore could reflect the operation of a memory system, in which the neuronal activity can be triggered by factors that probably include not only eye position command/feedback signals, but also probably vestibular and/or proprioceptive inputs. This representation of space "out there" would be an appropriate part of a primate memory system involved in memories of where in an environment an object was seen and more generally in the memory of particular events or episodes for which a spatial component normally provides part of the context.

Action Potentials↗

PubMed1998

Information about spatial view in an ensemble of primate hippocampal cells.

Hippocampal function was analyzed by making recordings from hippocampal neurons in monkeys actively walking in the laboratory. "Spatial view" cells, which respond when the monkey looks at a part of the environment, were analyzed. To assess quantitatively the information about the spatial environment represented by these cells, we applied information theoretic techniques to their responses. The average information provided by these cells about which location the monkey was looking at was 0.32 bits, and the mean across cells of the maximum information conveyed about which location was being looked at was 1.19 bits, measured in a period of 0.5 s. There were 16 locations for this analysis, each being one-quarter of one of the walls of the room. It also was shown that the mean spontaneous rate of firing of the neurons was 0.1 spikes/s, that the mean firing rate in the center of the spatial field of the neurons was 13.2 spikes/s, and that the mean sparseness of the representation measured in a 25-ms period was 0.04 and in a 500-ms time period was 0.19. (The sparseness is approximately equivalent to the proportion of the 25- or 500-ms periods in which the neurons showed one or more spikes.) Next it was shown that the mean size of the view fields of the neurons was 0.9 of a wall. In an approach to the issue of how an ensemble of neurons might together provide more precise information about spatial location than a single neuron, it was shown that in general the neurons had different centers for their view fields. It then was shown that the information from an ensemble of these cells about where in space is being looked at increases approximately linearly with the number of cells in the ensemble. This indicates that the number of places that can be represented increases approximately exponentially with the number of cells in the population. It is concluded that there is an accurate representation of space "out there" in the primate hippocampus. This representation of space out there would be an appropriate part of a primate memory system involved in memories of where in an environment an object was seen, and more generally in the memory of particular events or episodes, for which a spatial component normally provides part of the context.

Algorithms↗

PubMed1997

Responses of neurons in primary and inferior temporal visual cortices to natural scenes.

The primary visual cortex (V1) is the first cortical area to receive visual input, and inferior temporal (IT) areas are among the last along the ventral visual pathway. We recorded, in area V1 of anaesthetized cats and area IT of awake macaque monkeys, responses of neurons to videos of natural scenes. Responses were analysed to test various hypotheses concerning the nature of neural coding in these two regions. A variety of spike-train statistics were measured including spike-count distributions, interspike interval distributions, coefficients of variation, power spectra, Fano factors and different sparseness measures. All statistics showed non-Poisson characteristics and several revealed self-similarity of the spike trains. Spike-count distributions were approximately exponential in both visual areas for eight different videos and for counting windows ranging from 50 ms to 5 seconds. The results suggest that the neurons maximize their information carrying capacity while maintaining a fixed long-term-average firing rate, or equivalently, minimize their average firing rate for a fixed information carrying capacity.

Animals↗