Biomedical subjects

G L Gerstein

Publications and source records attributed to G L Gerstein.

At least 37 records · Page 2Linked to original sources

PubMed1996

Networks with lateral connectivity. I. dynamic properties mediated by the balance of intrinsic excitation and inhibition.

1. We studied the rapid dynamic changes of neuron response properties in the somatosensory cortex by the use of computer simulations. The model consists of three feedforward layers of spiking neurons, corresponding to skin, subcortex, and cortex structures. Measurements and analysis of model activity throughout this work are similar to those used in neurophysiological experiments. 2. The effects of various parameters on response properties of model neurons were investigated. The most important parameters were the lateral excitation and inhibition in the simulated cortical network. 3. The balance between excitation and inhibition is a key factor in determining the stability of the network model. There is a large excitation-inhibition (E-I) parameter region within which the model can stably respond to inputs. 4. The input-output relations and receptive field (RF) sizes of simulated neurons are modifiable by the E-I balance. The shapes of RFs are determined by both feedforward projections and the spatial distribution of lateral connections. 5. We simulated changes in temporal and spatial properties of neurons in response to manipulations that mimic bicuculine methiodide or glutamate application to the cortex. Simulation results agreed well with experimental data, suggesting that cortical transmitter levels play an important role in the dynamic responses of the neural net through their effects on E-I balance. 6. With parameters of the model set to an inhibition-dominant scheme, the model was able to reproduce experimentally observed rapid RF expansions that follow cortical lesion or input denervation. Simulation results also suggested that spontaneous inputs to a sensory system can serve as a source of tonic inhibition in the cortex. 7. We conclude that lateral connections could produce and maintain a cortical network having dynamic properties without the need to invoke synaptic plasticity. Individual neuron properties could be modified by changing the balance of cortical layer excitation and inhibition. In a real brain, this could be achieved either by changing levels of cortical transmitter (gamma-aminobutyric acid. for example) or by changing tonic background input to the cortical network.

Afferent Pathways↗

PubMed1996

Networks with lateral connectivity. II. Development of neuronal grouping and corresponding receptive field changes.

1. Using a three-layered network model defined in the previous paper, we studied the basic features of neurons in the cortical layer while the synaptic strengths of lateral excitatory connections were made modifiable by a Hebbian learning rule and a normalization process. 2. We found that neurons in the cortical layer formed groups through their lateral excitatory connections after the network was trained with sequential random dot stimulations. Neurons within a group connected tightly; neurons in different groups connected weakly. 3. The effects of model parameters and input parameters on the formation of neuronal groups were investigated. Results showed that the average size and rough shapes of groups were mainly determined by the spatial distribution of lateral connections within the cortical layer, irrespective of input parameters and training methods. Thus groups are structure dependent. 4. Lateral inhibition in the network is the only key factor that affects the grouping of neurons. Without an appropriate amount of distant inhibition, group formation does not occur. Group formation is very robust to all other parameters we tested. On the other hand, group locations are very easily disturbed by inputs or changes of parameters, suggesting that such neuronal groups are dynamically maintained. 5. With the development of neuronal groups, neurons can be divided into two response types. TN-1 neurons respond weakly to inputs and have small receptive fields or do not respond at all (silent); TN-II neurons, approximately 30-40% of all, respond strongly to inputs and have large receptive fields. The two types of neurons also differ with respect to response threshold and temporal firing patterns. After groups formed, receptive fields of TN-II neurons within the same group clustered spatially with high overlap, whereas receptive fields of TN-I neurons with detectable responses shifted systematically with the neuron's spatial location. 6. The two types of neurons are homogeneously distributed across the cortical layer. The population of each type of neuron produces a full representation of the input layer with weak or strong responses, respectively. 7. We concluded that neurons in the cortical network naturally assembled into functional groups. Such groups are dynamic and amenable to change by input stimuli. A fraction of neurons (30-40%) within the same group shares a similar receptive field and strongly respond together to stimuli, so that the network has more robust response to inputs. On the other hand, the responses of a large portion (60-70%) of neurons become weak or silent: these neurons are available for other (unknown) functional purposes.

Afferent Pathways↗

PubMed1996

Networks with lateral connectivity. III. Plasticity and reorganization of somatosensory cortex.

1. Mechanisms underlying cortical reorganizations were studied using a three-layered neural network model with neuronal groups already formed in the cortical layer. 2. Dynamic changes induced in cortex by behavioral training or intracortical microstimulation (ICMS) were simulated. Both manipulations resulted in reassembly of neuronal groups and formation of stimulus-dependent assemblies. Receptive fields of neurons and cortical representation of inputs also changed. Many neurons that had been weakly responsive or silent became active. 3. Several types of learning models were examined in simulating behavioral training, ICMS-induced dynamic changes, deafferentation, or cortical lesion. Each learning model most accurately reproduced features of experimental data from different manipulations, suggesting that more than one plasticity mechanism might be able to induce dynamic changes in cortex. 4. After skin or cortical stimulation ceased, as spontaneous activity continued, the stimulus-dependent assemblies gradually reverted into structure-dependent neuronal groups. However, relationships among individual neurons and identities of many neurons did not return to their original states. Thus a different set of neurons would be recruited by the same training stimulus sequence on its next presentation. 5. We also reproduced several typical long-term reorganizations caused by pathological manipulations such as cortical lesions, input loss, and digit fusion. 6. In summary, with Hebbian plasticity rules on lateral connections, the network model is capable of reproducing most characteristics of experiments on cortical reorganization. We propose that an important mechanism underlying cortical plastic changes is formation of temporary assemblies that are related to receipt of strongly synchronized localized input. Such stimulus-dependent assemblies can be dissolved by spontaneous activity after removal of the stimuli.

Afferent Pathways↗

PubMed1996

Nonlinearities within the cat LGN cell receptive fields in simulated network with recurrent inhibition.

We investigated the receptive fields of principal cells from the cat's lateral geniculate nucleus cells. About 20% of the X type neurones showed clear nonlinearities of summation when stimulated by two simultaneously onset, small bars of light. The possible source of this nonlinearity was studied on a specially designed model of a one-layer neuronal network with inhibitory, recurrent interactions, intended to mimic the inhibitory influence exerted on geniculate relay cells by perigeniculate interneurones. The model, when activated from periphery by two stimuli-like input patterns, produced at the output side the nonsymmetrical profiles of the receptive fields sensitivity, similar to those obtained in real experiments. This nonlinear output appeared when some of the relay cells were inhibited below their firing level threshold and this effect was spread through the network by lateral inhibitory connections. It is concluded that physiologically observed nonlinearities of the order of single receptive field mechanisms can be predicted by a simple recurrent network.

Animals↗

PubMed1994

Feature-linked synchronization of thalamic relay cell firing induced by feedback from the visual cortex.

The function of the massive feedback projection from visual cortex to its thalamic relay nucleus has so far eluded any clear overview. This feedback exerts a range of effects, including an increase in the inhibition elicited by moving contours, but the functional logic of the direct connections to the thalamic cells that relay the retinal input to the cortex remains largely unknown. In contrast to its thalamic nucleus, the visual cortex is characterized by cells that are strongly sensitive to the orientation of moving contours. Here we report that when driven by moving oriented visual stimuli the cortical feedback induces correlated firing in relay cells. This cortically induced correlation of relay cell activity produces coherent firing in those groups of relay cells with receptive field alignments appropriate to signalling the particular orientation of the moving contour to the cortex. Synchronization of relay cell firing means that they will elicit temporally overlapping excitatory postsynaptic potentials in their cortical target cells, thus increasing the chance that the cortical cells will fire. Effectively this increases the gain of the input for feature-linked events detected by the cortex. We propose that this feedback loop serves to lock or focus the appropriate circuitry onto the stimulus feature.

Action Potentials↗

PubMed1994

Rectification of correlation by a sigmoid nonlinearity.

We investigated the normalized autocovariance (correlation coefficient) function of the output of an erf() function nonlinearity subject to non-zero mean Gaussian noise input. When the sigmoid is wide compared to the input, or the input mean is close to the midpoint of the sigmoid, the output correlation coefficient function is very close to the input correlation coefficient function. When the noise mean and variance are such that there is a significant probability of operating in the saturation region and the sigmoid is not too flat, the correlation coefficient of the output function is less than that of the input. This difference is much greater when the correlation coefficient is negative than when it is positive. The sigmoid partially rectifies the correlation coefficient function. The analysis does not depend on the spectral properties of the input noise. All that is required is that the input at times t and (t + tau) be jointly gaussian with the same mean and autocovariance. The analysis therefore applies equally well to the case of two identical sigmoids with jointly gaussian inputs. This correlational rectification could help explain the parameter sensitivity of "neural network" models. If biological neurons share this property it could explain why few negative correlations between spike trains have been observed.

Animals↗

PubMed1994

Simulation of dynamic receptive fields in primary visual cortex.

A model network of spiking neurons with lateral connections was used to simulate short-term receptive field (RF) changes by removal of afferent input in the primary visual system. Several possible mechanisms for the dynamic RFs were explored and the simulation results were compared with experimental results obtained by Pettet and Gilbert [(1992) Proceedings of the National Academy of Science, U.S.A., 89, 8366-8370]. We found that appropriate input stimuli could induce a shift in the balance between modeled cortical lateral excitation and inhibition and in doing so cause RF expansion. Synaptic plasticity was neither necessary nor appropriate for short-term RF changes. An inhibition dominant network with neural adaptation successfully simulated Pettet and Gilbert's experiment of RF expansion and its reversibility induced by an "artificial scotoma". RF expansions induced by lesions were also explored with the model.

Action Potentials↗

PubMed1994

Neural ensemble coding in inferior temporal cortex.

1. Isolated, single-neuron extracellular potentials were recorded sequentially in area TE of the inferior temporal cortex (IT) of two macaque monkeys (n = 58 and n = 41 neurons). Data were obtained while the animals were performing a paired-associate task. The task utilized five stimuli and eight stimulus pairings (4 correct and 4 incorrect). Data were evaluated as average spike rate during experimental epochs of 100 or 400 ms. Single-unit and population characteristics were measured using a form of linear discriminant analysis and information theoretic measures. To evaluate the significance of covariance on population code measures, additional data consisting of simultaneous recordings from < or = 8 isolated neurons (n = 37) were obtained from a third macaque monkey that was passively viewing visual stimuli. 2. On average, 43% of IT neurons were activated by any of the stimuli used (60% if those inhibited also are included). Yet the neurons were rather unique in the relative magnitude of their responses to each stimulus in the test set. These results suggest that information may be represented in IT by the pattern of activity across neurons and that the representation is not sparsely coded. It is further suggested that the representation scheme may have similarities to DNA or computer codes wherein a coding element is not a local parametric descriptor. This is a departure from the V1 representation, which appears to be both local and parametric. It is also different from theories of IT representation that suggest a constructive basis set or "alphabet". From this view, determination of stimulus discrimination capacity in IT should be evaluated by measures of population activity patterns. 3. Evaluation of small groups of simultaneously recorded neurons obtained during a fixation task suggests that little information about visual stimuli is conveyed by covariance of activity in IT when a 100-ms time scale is used as in this study. This finding is consistent with a prior report, by Gochin et al., which used a 1-ms time scale and failed to find neural activity coherence or oscillations dependent on stimuli. 4. Population-stimulus-discrimination capacity measures were influenced by the number of neurons and to some extent the number and type of stimuli. 5. Information conveyed by individual neurons (mutual information) averaged 0.26 bits. The distribution of information values was unimodal and is therefore more consistent with a distributed than a local coding scheme.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

PubMed1993

A neural network model for texture discrimination.

A model of texture discrimination in visual cortex was built using a feedforward network with lateral interactions among relatively realistic spiking neural elements. The elements have various membrane currents, equilibrium potentials and time constants, with action potentials and synapses. The model is derived from the modified programs of MacGregor (1987). Gabor-like filters are applied to overlapping regions in the original image; the neural network with lateral excitatory and inhibitory interactions then compares and adjusts the Gabor amplitudes in order to produce the actual texture discrimination. Finally, a combination layer selects and groups various representations in the output of the network to form the final transformed image material. We show that both texture segmentation and detection of texture boundaries can be represented in the firing activity of such a network for a wide variety of synthetic to natural images. Performance details depend most strongly on the global balance of strengths of the excitatory and inhibitory lateral interconnections. The spatial distribution of lateral connective strengths has relatively little effect. Detailed temporal firing activities of single elements in the lateral connected network were examined under various stimulus conditions. Results show (as in area 17 of cortex) that a single element's response to image features local to its receptive field can be altered by changes in the global context.

Cybernetics↗

PubMed1992

Intrinsic signal optical imaging in the forepaw area of rat somatosensory cortex.

The responses of somatosensory cortex (S-I) to tactile stimulation of the forepaw were assessed by intrinsic signal optical imaging. The tips of digits two or five were alternately touched with mechanical tappers while video photographs were taken of S-I illuminated by an 800-nm light source. The resulting images showed two highlighted areas about 300 microns in diameter and 500 microns apart. Generation of these images required less than 1 hr. Electrode penetrations placed in the areas highlighted during stimulation provided multiunit recordings with receptive fields appropriate for the stimulated digit and not the other digit. Penetrations between the high-lighted areas yielded receptive fields on intervening digits. These results demonstrate that intrinsic signal optical images are obtainable in S-I and confirm the functional somatotopy previously reported using electrical recording. Furthermore, the short time required to produce the images and the obtainable spatial resolution suggest that optical recording could be employed for the study of cortical reorganization in this brain region.

Animals↗

PubMed1992

Respiratory-related neural assemblies in the brain stem midline.

1. The initial objective of this study was to determine whether respiratory-related neural assemblies exist in the brain stem midline. A second goal was to seek evidence for concurrent relationships among the neurons that could generate the detected synchrony. 2. Experiments were conducted on anesthetized, paralyzed, bilaterally vagotomized, artificially ventilated cats. Spike trains of four to nine simultaneously monitored neurons were recorded in the regions of n. raphe obscurus-n. raphe pallidus and n. raphe magnus. 3. Data were analyzed with cycle-triggered histograms, cross-correlograms, snowflakes, and the gravitational representation. A significance test for the gravity method was developed and tested with spike trains generated by simulated networks with defined connections. 4. Ninety-three groups of neurons from 24 cats were studied. Thirty-nine groups from 19 cats included neurons that discharged synchronously on a millisecond time scale; less than or equal to 19 pairs of synchronously discharging neurons were found in one group. Twenty-seven of these 39 groups included neurons that had respiratory-modulated firing rates and discharged synchronously with other group members. Synchronous assemblies included cells monitored at rostral or caudal locations, or both. 5. Six classes of relationships were inferred from groups of neurons with multiple correlations: divergence (n = 11); convergence (n = 7); connections with opposite actions between neurons (n = 5); projections of synchronous neurons to separate targets (n = 5); projections to one neuron in a synchronous group (n = 4); and projections between two synchronous groups with common elements (n = 6). 6. The results document the existence of assemblies of synchronously discharging respiratory-related neurons in midline regions of the brain stem and suggest that divergent excitatory and inhibitory connections within the midline participate in the generation of that synchrony. Links between assemblies may operate to stabilize their collective activity in a particular state.

Afferent Pathways↗

PubMed1992

Dynamic reconfiguration of brain stem neural assemblies: respiratory phase-dependent synchrony versus modulation of firing rates.

1. The objective of this work was to determine whether configurations of midline brain stem neural assemblies change during the respiratory cycle. 2. Spike trains of several single neurons were recorded simultaneously in anesthetized, paralyzed, bilaterally vagotomized, artificially ventilated cats. Data were analyzed with cross-correlational and gravity methods. 3. Sequential samples from each of eight groups of neurons known to contain synchronously discharging neurons exhibited temporal variations in that synchrony. 4. Gravity analysis of short (less than 200-s) samples of spike train data revealed 20 pairs of clustered particles that were not predicted from cross-correlation analysis of the parent data sets (greater than 20 min). 5. Twenty-nine groups of three to eight simultaneously monitored neurons, each with at least two synchronously discharging neurons, were analyzed for evidence of respiratory phase-dependent modulation of that coordinated activity. Spikes from successive interleaved inspiratory and expiratory intervals were analyzed separately. 6. Neurons pairs in 11 groups were more synchronous during the inspiratory interval; six groups had pairs that were more synchronous during the expiratory period. In two groups, different pairs were synchronous in different respiratory phases. In 11 of the 26 pairs that exhibited phase-dependent differences in synchrony, neither neuron had a respiratory-modulated firing rate as judged by either the cycle-triggered histogram or an analysis of variance of their firing rates. 7. Configurations of respiratory-related brain stem neural networks changed with time and the phases of breathing. Neurons with no apparent respiratory modulation of their individual firing rates collectively exhibited respiratory phase-dependent modulation of their impulse synchrony.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

PubMed1991

Underestimation of visual texture slant by human observers: a model.

The perspective image of an obliquely inclined textured surface exhibits shape and density distortions of texture elements which allow a human observer to estimate the inclination angle of the surface. However, it has been known since the work of Gibson (1950) that, in the absence of other cues, humans tend to underestimate the slant angle of the surface, particularly when the texture is perceived as being "irregular." The perspective distortions which affect texture elements also shift the projected spatial frequencies of the texture in systematic ways. Using a suitable local spectral filter to measure these frequency gradients, the inclination angle of the surface may be estimated. A computational model has been developed which performs this task using distributions of outputs from filters found to be a good description of simple-cell receptive fields. However, for "irregular" textures the filter output distributions are more like those of "regular" textures at shallower angles of slant, leading the computational algorithm to underestimate the slant angle. This behavioral similarity between human and algorithm suggests the possibility that a similar visual computation is performed in cortex.

Algorithms↗

PubMed1991

Cross-talk theory of memory capacity in neural networks.

The present paper presents a theory for the mechanics of cross-talk among constituent neurons in networks in which multiple memory traces have been embedded, and develops criteria for memory capacity based on the disruptive influences of this cross-talk. The theory is based on interconnection patterns defined by the sequential configuration model of dynamic firing patterns. The theory accurately predicts the memory capacities observed in computer simulated nets, and predicts that cortical-like modules should be able to store up to about 300-900 selectively retrievable memory traces before disruption by cross-talk is likely. It also predicts that the cortex may has designed itself for modules of 30,000 neurons to at least in part to optimize memory capacity.

Animals↗

PubMed1991

Functional interactions among neurons in inferior temporal cortex of the awake macaque.

Functional interactions among inferior temporal cortex (IT) neurons were studied in the awake, fixating macaque monkey during the presentation of visual stimuli. Extracellular recordings were obtained simultaneously from several microelectrodes, and in many cases, spike trains from more than one neuron were extracted from each electrode by the use of spike shape sorting technology. Functional interactions between pairs of neurons were measured using cross-correlation. Discharge patterns of single neurons were evaluated using auto-correlation and PST histograms. Neurons recorded on the same electrode (within about 100 microns) had more similar stimulus selectivity and were more likely to show functional interactions than those recorded on different electrodes spaced about 250 to 500 microns apart. Most neurons tended to fire in bursts tens to hundreds of milliseconds in duration, and asynchronously from the stimulus induced rate changes. Correlated neuronal firing indicative of shared inputs and direct interactions was observed. Occurrence of shared input was significantly lower for neuron pairs recorded on different electrodes than for neurons recorded on the same electrode. Direct connections occurred about as often for neurons on different electrodes as for neurons on the same electrode. These results suggest that input projections are usually restricted to less than 500 micron patches and are then distributed over greater distances by intrinsic connections. Measurements of synaptic contribution suggest that typically more than 5 near-simultaneous inputs are required to cause an IT neuron to discharge.

Animals↗

PubMed1990

Dynamic temporal properties of effective connections in rat dorsal cochlear nucleus.

In a prior report we presented evidence that functionally connected dorsal cochlear nucleus (DCN) neurons in close proximity can show differing peristimulus time histograms (PSTHs) in response to the same stimulus. We wished to further investigate how interconnections between such neurons might participate in the PSTH patterns. Methodology has recently been developed which permits measurement of rapid changes in effective connectivity between neuron pairs: the normalized joint PSTH. Using this technique we have observed that rapid changes in effective connectivity do occur in the DCN. These observations demonstrate that the effects of one DCN neuron on another cannot necessarily be understood by sequential recordings from single units, even if anatomical connectivity can be established.

Acoustic Stimulation↗

PubMed1989

Coordinated activity of neuron pairs in anesthetized rat dorsal cochlear nucleus.

We have recorded from small groups of neurons in the dorsal cochlear nucleus of anesthetized rats in an effort to study neuronal interactions. Multi-unit recordings on each single electrode were sorted by waveform into spike trains from individual neurons using a principal components spike sorter. Pairs of such sorted spike trains were studied with cross-correlation analysis to detect excitatory and/or inhibitory interactions. In a few cases recordings were obtained from two electrodes simultaneously, thus allowing cross-correlation studies without the consequences of spike train waveform sorting. All neurons were characterized by their strongest response frequency (at a fixed sound pressure level) and peristimulus histogram responses to 55 ms tone bursts. Fifty-eight percent of the neuron pairs studied showed peaks in their cross-correlograms indicative of coordinated neural activity. Of these pairs, 86% showed peak configurations (i.e. correlograms with asymmetrically located peaks) consistent with the interpretation that one cell induced the other to discharge. The remaining correlograms contained symmetric peaks which were centrally located, possibly due to shared input to these neuron pairs. Latencies of asymmetric peaks in cross-correlograms were typically 2 ms; consequently, an intervening excitatory synapse may be involved. Similar results were obtained from at least one pair of neurons where each neuron was recorded by a separate electrode. Strongest response frequencies of each neuron pair, for which they could be determined, were within 0.17 log units. Peristimulus histograms from each neuron in these pairs revealed that it was common for adjacent cells to respond with differing time patterns under the same stimulus conditions. The variations in histogram patterns of interconnected neurons suggests some relatively complex integrative function for these circuits.

Acoustic Stimulation↗

PubMed1989

Gravitational representation of simultaneously recorded brainstem respiratory neuron spike trains.

Experiments designed to study concurrent processes in neural networks have been hampered by limitations of available analytical methods. A recently described gravitational representation of spike train data was used to evaluate groups of simultaneously monitored medullary respiratory related neurons in anesthetized, vagotomized cats. The results establish that the method can detect and define functional associations among elements of such groups after as few as 20 respiratory cycles.

Action Potentials↗