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E J Chichilnisky

Publications and source records attributed to E J Chichilnisky.

At least 19 recordsLinked to original sources
PubMed2006

The structure of multi-neuron firing patterns in primate retina.

Current understanding of many neural circuits is limited by our ability to explore the vast number of potential interactions between different cells. We present a new approach that dramatically reduces the complexity of this problem. Large-scale multi-electrode recordings were used to measure electrical activity in nearly complete, regularly spaced mosaics of several hundred ON and OFF parasol retinal ganglion cells in macaque monkey retina. Parasol cells exhibited substantial pairwise correlations, as has been observed in other species, indicating functional connectivity. However, pairwise measurements alone are insufficient to determine the prevalence of multi-neuron firing patterns, which would be predicted from widely diverging common inputs and have been hypothesized to convey distinct visual messages to the brain. The number of possible multi-neuron firing patterns is far too large to study exhaustively, but this problem may be circumvented if two simple rules of connectivity can be established: (1) multi-cell firing patterns arise from multiple pairwise interactions, and (2) interactions are limited to adjacent cells in the mosaic. Using maximum entropy methods from statistical mechanics, we show that pairwise and adjacent interactions accurately accounted for the structure and prevalence of multi-neuron firing patterns, explaining approximately 98% of the departures from statistical independence in parasol cells and approximately 99% of the departures that were reproducible in repeated measurements. This approach provides a way to define limits on the complexity of network interactions and thus may be relevant for probing the function of many neural circuits.

Action Potentials↗
PubMed2006

Electrical stimulation of mammalian retinal ganglion cells with multielectrode arrays.

Existing epiretinal implants for the blind are designed to electrically stimulate large groups of surviving retinal neurons using a small number of electrodes with diameters of several hundred micrometers. To increase the spatial resolution of artificial sight, electrodes much smaller than those currently in use are desirable. In this study, we stimulated and recorded ganglion cells in isolated pieces of rat, guinea pig, and monkey retina. We used microfabricated hexagonal arrays of 61 platinum disk electrodes with diameters between 6 and 25 microm, spaced 60 microm apart. Charge-balanced current pulses evoked one or two spikes at latencies as short as 0.2 ms, and typically only one or a few recorded ganglion cells were stimulated. Application of several synaptic blockers did not abolish the evoked responses, implying direct activation of ganglion cells. Threshold charge densities were typically <0.1 mC/cm2 for a pulse duration of 100 micros, corresponding to charge thresholds of <100 pC. Stimulation remained effective after several hours and at high frequencies. To show that closely spaced electrodes can elicit independent ganglion cell responses, we used the multielectrode array to stimulate several nearby ganglion cells simultaneously. From these data, we conclude that electrical stimulation of mammalian retina with small-diameter electrode arrays is achievable and can provide high temporal and spatial precision at low charge densities. We review previous epiretinal stimulation studies and discuss our results in the context of 32 other publications, comparing threshold parameters and safety limits.

Action Potentials↗
PubMed2005

Prediction and decoding of retinal ganglion cell responses with a probabilistic spiking model.

Sensory encoding in spiking neurons depends on both the integration of sensory inputs and the intrinsic dynamics and variability of spike generation. We show that the stimulus selectivity, reliability, and timing precision of primate retinal ganglion cell (RGC) light responses can be reproduced accurately with a simple model consisting of a leaky integrate-and-fire spike generator driven by a linearly filtered stimulus, a postspike current, and a Gaussian noise current. We fit model parameters for individual RGCs by maximizing the likelihood of observed spike responses to a stochastic visual stimulus. Although compact, the fitted model predicts the detailed time structure of responses to novel stimuli, accurately capturing the interaction between the spiking history and sensory stimulus selectivity. The model also accounts for the variability in responses to repeated stimuli, even when fit to data from a single (nonrepeating) stimulus sequence. Finally, the model can be used to derive an explicit, maximum-likelihood decoding rule for neural spike trains, thus providing a tool for assessing the limitations that spiking variability imposes on sensory performance.

Action Potentials↗
PubMed2005

Detection sensitivity and temporal resolution of visual signals near absolute threshold in the salamander retina.

Several studies have suggested that the visual system can detect dim lights with a fidelity limited only by Poisson fluctuations in photon absorption and spontaneous activation of rhodopsin. If correct, this implies that neural processing of responses produced by rod photoreceptors is efficient and effectively noiseless. However, experimental uncertainty makes this conclusion tenuous. Furthermore, previous work provided no information about how accurately stimulus timing is represented. Here, the detection sensitivity and temporal resolution of salamander rods and retinal ganglion cells (RGCs) are compared in nearly matched experimental conditions by using recorded responses to identify the time of a flash. At detection threshold, RGCs could reliably signal the absorption of 20-50 photons, but the rods within the RGC receptive field could signal stimuli 3-10 times weaker. For flash strengths 10 times higher than detection threshold, some RGCs could distinguish stimulus timing with a resolution finer than 100 msec, within a factor of 2 of the rod limit. The relationship between RGC and rod sensitivity could not be explained by added noise in the retinal circuitry but could be explained by a threshold acting after pooling of rod signals. Simulations of rod signals indicated that continuous noise, rather than spontaneous activation of rhodopsin or fluctuations in the single-photon response, limited temporal resolution. Thus, detection of dim lights was limited by retinal processing, but, at higher light levels, synaptic transmission, cellular integration of synaptic inputs, and spike generation in RGCs faithfully conveyed information about the time of photon absorption.

Ambystoma↗
PubMed2005

Estimating entropy rates with Bayesian confidence intervals.

The entropy rate quantifies the amount of uncertainty or disorder produced by any dynamical system. In a spiking neuron, this uncertainty translates into the amount of information potentially encoded and thus the subject of intense theoretical and experimental investigation. Estimating this quantity in observed, experimental data is difficult and requires a judicious selection of probabilistic models, balancing between two opposing biases. We use a model weighting principle originally developed for lossless data compression, following the minimum description length principle. This weighting yields a direct estimator of the entropy rate, which, compared to existing methods, exhibits significantly less bias and converges faster in simulation. With Monte Carlo techinques, we estimate a Bayesian confidence interval for the entropy rate. In related work, we apply these ideas to estimate the information rates between sensory stimuli and neural responses in experimental data (Shlens, Kennel, Abarbanel, & Chichilnisky, in preparation).

Bayes Theorem↗
PubMed2004

Fidelity of the ensemble code for visual motion in primate retina.

Sensory experience typically depends on the ensemble activity of hundreds or thousands of neurons, but little is known about how populations of neurons faithfully encode behaviorally important sensory information. We examined how precisely speed of movement is encoded in the population activity of magnocellular-projecting parasol retinal ganglion cells (RGCs) in macaque monkey retina. Multi-electrode recordings were used to measure the activity of approximately 100 parasol RGCs simultaneously in isolated retinas stimulated with moving bars. To examine how faithfully the retina signals motion, stimulus speed was estimated directly from recorded RGC responses using an optimized algorithm that resembles models of motion sensing in the brain. RGC population activity encoded speed with a precision of approximately 1%. The elementary motion signal was conveyed in approximately 10 ms, comparable to the interspike interval. Temporal structure in spike trains provided more precise speed estimates than time-varying firing rates. Correlated activity between RGCs had little effect on speed estimates. The spatial dispersion of RGC receptive fields along the axis of motion influenced speed estimates more strongly than along the orthogonal direction, as predicted by a simple model based on RGC response time variability and optimal pooling. on and off cells encoded speed with similar and statistically independent variability. Simulation of downstream speed estimation using populations of speed-tuned units showed that peak (winner take all) readout provided more precise speed estimates than centroid (vector average) readout. These findings reveal how faithfully the retinal population code conveys information about stimulus speed and the consequences for motion sensing in the brain.

Action Potentials↗
PubMed2004

Blue-yellow signals are enhanced by spatiotemporal luminance contrast in macaque V1.

We measured the color tuning of a population of S-cone-driven V1 neurons in awake, fixating monkeys. Analysis of randomly chosen color stimuli that were effective in evoking action potentials showed that these neurons received opposite sign input from the S cones and a combination of L and M cones. Surprisingly, these cells also responded to LM cone contrast irrespective of polarity, a nonlinear sensitivity that was masked by conventional linear analysis methods. Taken together, these observations can be summarized in a nonlinear model that combines nonopponent and opponent signals such that luminance contrast enhances color processing. These findings indicate that important aspects of the cortical representation of color cannot be described by classical linear analysis, and reveal a possible neural correlate of perceptual color-luminance interactions.

Action Potentials↗
PubMed2004

Precision of spike trains in primate retinal ganglion cells.

Recent studies have revealed striking precision in the spike trains of retinal ganglion cells in several species and suggested that this precision could be an important aspect of visual signaling. However, the precision of spike trains has not yet been described in primate retina. The spike time and count variability of parasol (magnocellular-projecting) retinal ganglion cells was examined in isolated macaque monkey retinas stimulated with repeated presentations of high contrast, spatially uniform intensity modulation. At the onset of clearly delineated periods of firing, retinal ganglion cells fired spikes time-locked to the stimulus with a variability across trials as low as 1 ms. Spike count variance across trials was much lower than the mean and sometimes approached the minimum variance possible with discrete counts, inconsistent with Poisson statistics expected from independently generated spikes. Spike time and count variability decreased systematically with stimulus strength. These findings were consistent with a model in which firing probability was determined by a stimulus-driven free firing rate modulated by a recovery function representing the action potential absolute and relative refractory period.

Action Potentials↗
PubMed2003

Temporal resolution of ensemble visual motion signals in primate retina.

Recent studies have examined the temporal precision of spiking in visual system neurons, but less is known about the time scale that is relevant for behaviorally important visual computations. We examined how spatiotemporal patterns of spikes in ensembles of primate retinal ganglion cells convey information about visual motion to the brain. The direction of motion of a bar was estimated by comparing the timing of responses in ensembles of parasol (magnocellular-projecting) retinal ganglion cells recorded simultaneously, using a cross-correlation approach similar to standard models of motion sensing. To identify the temporal resolution of motion signals, spike trains were low-pass filtered before estimating the direction of motion. The filter time constant that resulted in most accurate motion sensing was in the range of 10-50 msec for a range of stimulus speeds and contrasts and approached a lower limit of approximately 10 msec at high speeds and contrasts. This time constant was, on average, comparable to the length of interspike intervals. These findings suggest that cortical neurons could filter their inputs on a time scale of tens of milliseconds, rather than relying on the precise times of individual input spikes, to sense motion most reliably.

Action Potentials↗
PubMed2002

Functional asymmetries in ON and OFF ganglion cells of primate retina.

Functional asymmetries in the ON and OFF pathways of the primate visual system were examined using simultaneous multi-electrode recordings from dozens of retinal ganglion cells (RGCs) in vitro. Light responses of RGCs were characterized using white noise stimulation. Two distinct functional types of cells frequently encountered, one ON and one OFF, had non-opponent spectral sensitivity, relatively high response gain, transient light responses, and large receptive fields (RFs) that tiled the region of retina recorded, suggesting that they belonged to the same morphological cell class, most likely parasol. Three principal functional asymmetries were observed. (1) Receptive fields of ON cells were 20% larger in diameter than those of OFF cells, resulting in higher full-field sensitivity. (2) ON cells had faster response kinetics than OFF cells, with a 10-20% shorter time to peak, trough and zero crossing in the biphasic temporal impulse response. (3) ON cells had more nearly linear light responses and were capable of signaling decrements, whereas OFF cells had more strongly rectifying responses that provided little information about increments. These findings suggest specific mechanistic asymmetries in retinal ON and OFF circuits and differences in visual performance on the basis of the activity of ON and OFF parasol cells.

Action Potentials↗
PubMed2001

Adaptation to temporal contrast in primate and salamander retina.

Visual adaptation to temporal contrast (intensity modulation of a spatially uniform, randomly flickering stimulus) was examined in simultaneously recorded ensembles of retinal ganglion cells (RGCs) in tiger salamander and macaque monkey retina. Slow contrast adaptation similar to that recently discovered in salamander and rabbit retina was observed in monkey retina. A novel method was developed to quantify the effect of temporal contrast on steady-state sensitivity and kinetics of light responses, separately from nonlinearities that would otherwise significantly contaminate estimates of sensitivity. Increases in stimulus contrast progressively and reversibly attenuated and sped light responses in both salamander and monkey RGCs, indicating that a portion of the contrast adaptation observed in visual cortex originates in the retina. The effect of adaptation on sensitivity and kinetics differed in simultaneously recorded populations of ON and OFF cells. In salamander, adaptation affected the sensitivity of OFF cells more than ON cells. In monkey, adaptation affected the sensitivity of ON cells more than OFF cells. In both species, adaptation sped the light responses of OFF cells more than ON cells. Functionally defined subclasses of ON and OFF cells also exhibited asymmetric adaptation. These findings indicate that contrast adaptation differs in parallel retinal circuits that convey distinct visual signals to the brain.

Action Potentials↗
PubMed2001

A simple white noise analysis of neuronal light responses.

A white noise technique is presented for estimating the response properties of spiking visual system neurons. The technique is simple, robust, efficient and well suited to simultaneous recordings from multiple neurons. It provides a complete and easily interpretable model of light responses even for neurons that display a common form of response nonlinearity that precludes classical linear systems analysis. A theoretical justification of the technique is presented that relies only on elementary linear algebra and statistics. Implementation is described with examples. The technique and the underlying model of neural responses are validated using recordings from retinal ganglion cells, and in principle are applicable to other neurons. Advantages and disadvantages of the technique relative to classical approaches are discussed.

Action Potentials↗
PubMed1999

Trichromatic opponent color classification.

Stimuli varying in intensity and chromaticity, presented on numerous backgrounds, were classified into red/green, blue/yellow and white/black opponent color categories. These measurements revealed the shapes of the boundaries that separate opponent colors in three-dimensional color space. Opponent color classification boundaries were generally not planar, but their shapes could be summarized by a piecewise linear model in which increment and decrement color signals are combined with different weights at two stages to produce opponent color sensations. The effect of background light on classification was largely explained by separate gain changes in increment and decrement cone signals.

Adaptation, Ocular↗
PubMed1999

Receptive-field microstructure of blue-yellow ganglion cells in primate retina.

We examined the functional microcircuitry of cone inputs to blue-ON/yellow-OFF (BY) ganglion cells in the macaque retina using multielectrode recording. BY cells were identified by their ON responses to blue light and OFF responses to red or green light. Cone-isolating stimulation indicated that ON responses originated in short (S) wavelength-sensitive cones, whereas OFF responses originated in both long (L) and middle (M) wavelength-sensitive cones. Stimulation with fine spatial patterns revealed locations of individual S cones in BY cell receptive fields. Neighboring BY cells received common but unequal inputs from one or more S cones. Inputs from individual S cones differed in strength, indicating different synaptic weights, and summed approximately linearly to control BY cell firing.

Animals↗
PubMed1995

Photoreceptor sensitivity changes explain color appearance shifts induced by large uniform backgrounds in dichoptic matching.

Photoreceptor sensitivity changes explained the effect of large uniform backgrounds on the color appearance of small targets in a dichoptic asymmetric color matching experiment. Subjects viewed in each eye a target superimposed on a large background. The backgrounds presented to the two eyes had different spectral compositions. Subjects adjusted the target seen by the right eye to match the appearance of the target seen by the left eye. Receptor sensitivity changes explained the effect of numerous adapting backgrounds on the color appearance of many targets with high precision. Post-receptoral sensitivity changes provided a poorer account of the data. The apparent sensitivity of each receptor class varied inversely with changes in background light absorbed by that receptor class, but did not depend on background light absorbed by the other two receptor classes.

Adaptation, Ocular↗
PubMed1993

Functional segregation of color and motion perception examined in motion nulling.

We examine two hypotheses about the functional segregation of color and motion perception, using a motion nulling task. The most common interpretation of functional segregation, that motion perception depends only on one of the three dimensions of color, is rejected. We propose and test an alternative formulation of functional segregation: that motion perception depends on a univariate motion signal driven by all three color dimensions, and that the motion signal is determined by the product of the stimulus contrast and a term that depends only on the relative cone excitations. Two predictions of this model are confirmed. First, motion nulling is transitive: when two stimuli null a third they also null another. Second, motion nulling is homogeneous: if two stimuli null one another, they continue to null one another when their contrasts are scaled equally. We describe how to apply our formulation of functional segregation to other behavioral and physiological measurements.

Color Perception↗
PubMed1986

A mathematical model of pattern formation.

This paper presents an explicit mathematical model describing pattern formation in monolayer epithelia. The approach is a generalization of the equations describing soap bubble configurations (Plateau, 1873; Thompson, 1917; Almgren & Taylor, 1976) that allows adjacent cells to adhere with differing intensities (Steinberg, 1962, 1978). The model is a system of simultaneous non-linear equations that considers cell-cell interactions in a two-dimensional sheet. The implementation involves using the equations of the model to predict explicitly the energy-minimizing configuration of a system of cells, based on the adhesivity of their membranes. The model can thus be used to explore the effects of varying adhesions on the dynamics of pattern formation. Following Chichilnisky (1985), such a descriptive system is introduced in this paper, and its predictive properties explored.

Cell Aggregation↗