Synaptic organization and ionic basis of on and off channels in mudpuppy retina. II. Chloride-dependent ganglion cell mechanisms

Extracellular ganglion cell recordings in the perfused mudpuppy eyecup show that a chloride-free (c-f) perfusate abolishes the center and surround excitation of on-center cells, the surround excitation of off- center cells, and the on discharge of on-off cells. These changes in ganglion cell receptive field organization are anticipated in view of the effects of a c-f environment on the neurons which are presynaptic to the ganglion cells. However, chloride-dependent inhibitory postsynaptic (IPS) responses have been observed in on-off ganglion cells. These inhibitory postsynaptic potentials (IPSP's) are preceeded by (ESPS's) exitatory postsynaptic potentials and are apparently mediated by amacrine cells. The light-activated hyperpolarization of off cells is not the result of a chloride-dependent IPSP and probably results from disfacilitation.     

A spatio-temporal model of ganglion cell receptive field in the cat retina

A spatio-temporal model of ganglion cell receptive fields is proposed on the basis of receptive field characteristics of cat retinal ganglion cells reported in our previous paper. The model consists of the linear and nonlinear mechanisms in the ganglion cell receptive field. The linear mechanism is assumed to be composed of antagonistic center and surround mechanisms. Then, by integrating these mechanisms we construct a spatio-temporal impulse response function of ganglion cell receptive field. Here we assume that spatio-temporal impulse response function may be factored into spatial and temporal terms. By Fouriertransforming the spatio-temporal impulse response function, we can obtain the spatio-temporal transfer function. Contrast sensitivity characteristics of X-and Y-cells in the cat retina may be explained by the transfer function.     

Synaptic organization and ionic basis of on and off channels in mudpuppy retina. I. Intracellular analysis of chloride-sensitive electrogenic properties of receptors, horizontal cells, bipolar cells, and amacrine cells

Intracellular recordings from receptors, horizontal cells, bipolars, and amacrines have been carried out in the perfused mudpuppy eyecup. The introduction of a chloride-free (c-f) medium results in initial transient potential changes in many cells followed by a slow loss of light-evoked activity of the depolarizing bipolar, the horizontal cell, and the on depolarization of amacrine cells. The hyperpolarizing bipolar remains responsive to light stimulation in a c-f medium, but the antagonistic surround mechanism is abolished. These effects are reversible after returning to a normal ionic medium. The results of this study provide insight into the retinal connections which underlie ganglion cell receptive field organization. It is concluded that the depolarizing bipolar is excitatory to on ganglion cells and is also the pathway for on-excitation of on-off cells. The hyperpolarizing bipolar mediates the off discharge of off and on-off cells. Amacrine cells receive input from both depolarizing and hyperpolarizing bipolar cells. These findings raise the possibility that transmembrane movements of chloride ions are critical for the light responsiveness of horizontal and depolarizing bipolar cell activity.     

Influence of amacrine cells on receptive field organization of ganglion cells of the generalized vertebrate cone retina: Electronic simulation

Two classes of amacrine cells are simulated, small-field and large-field. Small-field amacrine cells are formed by input from a single bipolar cell, while large-field amacrine cell is formed by inputs from same 7 bipolar cells that form the ganglion cell. Only tonic amacrine cells are studied with both chromatic and luminosity types as well as double-and single-opponent receptive fields. Amacrine cells are used in both feedforward to ganglion cells and feedback to bipolar and horizontal cells. Feedback to bipolar cells or feedfoward to ganglion cells affected steady state levels in a predictable fashion. Negative feedback to bipolar cells and positive feedfoward to ganglion cells does not introduce transients to ganglion cells while negative feedback to horizontal cells and negative feedfoward does. Feedback to horizontal cells produces complex effects on bipolar, amacrine and ganglion cells dependent on such factors as center-surround field balance and negative feedback from luminosity type of horizontal cell to cones.     

A dynamic model of the receptive field of L-cells in the carp retina

A dynamic model of the receptive field of L2-cells in the carp retina is developed by using our experimental results on the basis of physiological and morphological evidences. Linear spatial summation is assumed in the model for the interactions among L2-cells. Linear forward and feedback loops are also assumed for the interactions between L2-cells and cones. The model has dynamic properties similar to the ones of the receptive field of L2-cells: L2-cells respond faster as the size of a light spot is enlarged and the L2-cells nearer to the center of the light spot respond faster. It is suggested that the faster responding properties of L2-cells are due to the feedback action.     

A model for the temporal organization of X- and Y-type receptive fields in the primate retina

A model is proposed for the temporal characteristics of X-and Y-type responses of ganglion cells in the primate retina. The main suggestions of the model are: (I) The X-type temporal response is determined primarily by the delay between center and surround contributions. (II) The Y-type response is generated in the inner plexiform layer by a derivativelike operation on the bipolar cell's input, followed by a rectification in the convergence of these inputs onto the Y-ganglion-cell. (III) The derivative-like operation is obtained by recurrent inhibition in the dyad synaptic structure.The X-and Y-type responses predicted by the model, for a variety of stimuli, were examined and compared with available electrophysiological recordings. Finally, certain predictions derived from the model are discussed.     

A model of anuran retina relating interneurons to ganglion cell responses

A model is presented which accounts for many characteristic response properties used to classify anuran ganglion cell types while being consistent with data concerning interneurons. In the model color is ignored and input stimuli are assumed to be only black and white at high contrast. We show that accurate ganglion cell responses are obtained even with simplified receptors and horizontal cells: Receptors are modeled as responding with a step change, while horizontal cells respond only to global changes in intensity brought about by full field illumination changes. A hyperpolarizing and depolarizing bipolar cell are generated y subtracting local receptor and horizontal potentials. Two transient amacrine cells (On and Off) are generated using a high-pass filter like mechanism with a thresholded output which responds to positive going changes in the corresponding bipolar cell potentials. The model shows how a selective combination of bipolar and amacrine channels can account for many of the response properties used to classify the anuran ganglion cell types (class-0 through 4) and makes several experimental predictions.     

The involvement of gamma-aminobutyric acid in the organization of cat retinal ganglion cell receptive fields. A study with picrotoxin and bicuculline

The effects of picrotoxin and bicuculline upon the discharge pattern of center-surround organized cat retinal ganglion cells of X and Y type were studied. All experiments were carried out under scotopic or possibly low mesopic conditions; mostly but not exclusively on-center cells were studied. Stimuli were chosen so that responses were either; (a) "purely" central; (b) surround dominated; or (c) clearly mixed but center dominated. In each case a pre-drug control response was estaboished, the drug was administered intravenously, and its subsequent effect upon the response was observed. In Y cells both picrotoxin and bicucullin caused the center-driven component of the response to become somewhat reduced in magnitude, while the surround component was substantially reduced. There was thus a change in center- surround balance in favor of the center-driven component. Responses of X cells remained virtually unaffected by both picrotoxin and bicuculline.     

A possible mechanism of stochastic resonance in the light of an extra-classical receptive field model of retinal ganglion cells

Traditionally the intensity discontinuities in an image are detected as zero-crossings of the second derivative with the help of a Laplacian of Gaussian (LOG) operator that models the receptive field of retinal Ganglion cells. Such zero-crossings supposedly form a raw primal sketch edge map of the external world in the primary visual cortex of the brain. Based on a new operator which is a linear combination of the LOG and a Dirac-delta function that models the extra-classical receptive field of the ganglion cells, we find that zero-crossing points thus generated, store in presence of noise, apart from the edge information, the shading information of the image in the form of density variation of these points. We have also shown that an optimal image contrast produces best mapping of the shading information to such zero-crossing density variation for a given amount of noise contamination. Furthermore, we have observed that an optimal amount of noise contamination reproduces the minimum optimal contrast and hence gives rise to the best representation of the original image. We show that this phenomenon is similar in nature to that of stochastic resonance phenomenon observed in psychophysical experiments.     

A cooperation and competition based simple cell receptive field model and study of feed-forward linear and nonlinear contributions to orientation selectivity

We present a model for development of orientation selectivity in layer IV simple cells. Receptive field (RF) development in the model, is determined by diffusive cooperation and resource limited competition guided axonal growth and retraction in geniculocortical pathway. The simulated cortical RFs resemble experimental RFs. The receptive field model is incorporated in a three-layer visual pathway model consisting of retina, LGN and cortex. We have studied the effect of activity dependent synaptic scaling on orientation tuning of cortical cells. The mean value of hwhh (half width at half the height of maximum response) in simulated cortical cells is 58° when we consider only the linear excitatory contribution from LGN. We observe a mean improvement of 22.8° in tuning response due to the non-linear spiking mechanisms that include effects of threshold voltage and synaptic scaling factor.     

Hydrodynamic model of temperature change in open ionic channels.

Most theories of open ionic channels ignore heat generated by current flow, but that heat is known to be significant when analogous currents flow in semiconductors, so a generalization of the Poisson-Nernst-Planck theory of channels, called the hydrodynamic model, is needed. The hydrodynamic theory is a combination of the Poisson and Euler field equations of electrostatics and fluid dynamics, conservation laws that describe diffusive and convective flow of mass, heat, and charge (i.e., current), and their coupling. That is to say, it is a kinetic theory of solute and solvent flow, allowing heat and current flow as well, taking into account density changes, temperature changes, and electrical potential gradients. We integrate the equations with an essentially nonoscillatory shock-capturing numerical scheme previously shown to be stable and accurate. Our calculations show that 1) a significant amount of electrical energy is exchanged with the permeating ions; 2) the local temperature of the ions rises some tens of degrees, and this temperature rise significantly alters for ionic flux in a channel 25 A long, such as gramicidin-A; and 3) a critical parameter, called the saturation velocity, determines whether ionic motion is overdamped (Poisson-Nernst-Planck theory), is an intermediate regime (called the adiabatic approximation in semiconductor theory), or is altogether unrestricted (requiring the full hydrodynamic model). It seems that significant temperature changes are likely to accompany current flow in the open ionic channel.     

Stimulus size selectivity and receptive field organization of ectostriatal neurons in the pigeon

The avian ectostriatum is the telencephalic recipient zone of the tectofugal pathway, which may be homologous to the colliculo-pulvinar-cortical pathway in mammals. The present paper studies the visual response properties and receptive field organization of ectostriatal cells in pigeons with extracellular recording and computer mapping techniques. The results indicate that 90% of ectostriatal cells prefer forward, downward and upward motion of visual stimuli at velocities in the range of 16-128 degrees s(-1). They respond optimally to small stimuli (1-4 degrees visual angle), mostly preferring a target of 2 degrees. Most cells (78.8%) have one excitatory receptive field that usually possesses one or two hotspots, some cells (13.5%) have two excitatory receptive fields each having one or two hotspots, and a few cells (7.7%) have one excitatory receptive field with an unresponsive region in the center. An inhibitory receptive field is not found surrounding the excitatory receptive field in the ectostriatal cells examined. These response properties and receptive field organization may reflect the possible roles of the ectostriatum in stimulus discrimination and visual cognition.     

Thresholds and latencies of retinal ganglion cell responses in cats to local stimulation of various parts of their receptive field

Depending on their responses to separate stimulation of the center and periphery of the receptive field, all ganglion cells of the cat retina can be subdivided into two types: ON-center (OFF-periphery) and OFF-center (ON-periphery). By all the parameters studied these ON- and OFF-systems were symmetrical. This apparently reflects, first, the equality of informativeness of illumination and darkening of individual areas of the visual field and, second, adaptation in order to widen the dynamic range of the visual channel of information transmission. Thresholds of unit responses to stimulation of the periphery and center of their receptive fields were identical. The latent periods of the unit responses were much longer in the first case than in the second. This is regarded as providing the functional basis for discrimination between "central" and "peripheral" unit responses by higher structures.Institute of Control Problems, Academy of Sciences of the USSR. Translated from Neirofiziologiya, Vol. 3, No. 6, pp. 644–649, November–December, 1971.     

A domain model for eukaryotic DNA organization: a molecular basis for cell differentiation and chromosome evolution

A model for eukaryotic chromatin organization is presented in which the basic structural and functional unit is the DNA domain. This simple model predicts that both chromosome replication and cell type-specific control of gene expression depend on a combination of stable and dynamic DNA-nuclear matrix interactions. The model suggests that in eukaryotes, DNA regulatory processes are controlled mainly by the intranuclear compartmentalization of the specific DNA sequences, and that control of gene expression involves multiple steps of specific DNA-nuclear matrix interactions. Predictions of the model are tested using available biochemical, molecular and cell biological data. In addition, the domain model is discussed as a simple molecular mechanism to explain cell differentiation in multi-cellular organisms and to explain the evolution of eukaryotic genomes consisting mainly of repetitive sequences and "junk" DNA.     

Dissociation of field potential from neuronal activity in the isolated retina: failure of the b-wave with normal ganglion cell response.

The b-wave of the isolated rabbit retina was compared with the ganglion cell response to light before and after modification of the retina's incubating medium. Marked diminution of the b-wave, with no reduction in ganglion cell response, was observed under three experimental conditions: (1) following a short period of anoxia; (2) following a short period in 0.2 mM Ca++; (3) in a small percentage of preparations, simply as a result of prolonged incubation in control medium. In contrast, a short period in 50 mM K+ led to a parallel fall and parallel recovery of both responses. It is apparent that under selected conditions the field potentials which constitute the b-wave are poorly correlated with the retina's neural activity.     

Activity sharpens the regenerating retinotectal projection in goldfish: sensitive period for strobe illumination and lack of effect on synaptogenesis and on ganglion cell receptive field properties

The regenerating optic nerve of goldfish first reestablishes a rough retinotopic map on the contralateral tectum and then sharpens it. Disruption of visual activity, either by blocking activity with intraocular tetrodotoxin (TTX; Schmidt and Edwards, 1983) or by synchronizing activity with strobe illumination (Schmidt and Eisele, 1985), disrupts the sharpening process: the map is correctly oriented but the multiunit receptive fields at each point average 25-40 degrees in diameter. In order to test whether strobe and TTX interfere with the same mechanism, we have tested whether their sensitive periods are the same, and whether strobe, like TTX treatment, does not affect either ganglion cell receptive field properties or synaptogenesis. In parallel studies, we exposed fish to 2 weeks of either strobe illumination or intraocular TTX beginning at various times after crush and determined via electrophysiological recordings that the periods of sensitivity were nearly identical. There was no effect of either treatment during the first 2 weeks (before the fibers arrive at the tectum), maximal disruption of sharpening between 14 and 50 days (the period of rapid synaptogenesis), decreasing disruption between 50 and 125 days, and no effect beyond that point or in the normal projection. In addition, long strobe exposures of up to 142 days produced no greater disruptions than shorter 2-3-week exposures, indicating no cumulative effect. The reestablishment of synaptic transmission in tectum, assayed by recording field potentials elicited by optic nerve shock, was not affected by stroboscopic illumination. Finally, individual ganglion cells, recorded intraretinally following long-term strobe exposure, had receptive fields that were normal both in size and in their characteristic responses to light-on, to light-off, or to both on and off. These findings support the hypothesis that strobe-like TTX prevents retinotopic refinement by preventing the correction of errors initially made by the ingrowing optic axons (Schmidt et al., 1988).     

Self organization of the microtubule network. A diffusion based model

Microtubules form organized polymer networks in cells. Experimental evidence indicates that their mechanical properties do not play a significant role in the generation of such regular patterns. This spatial organization seems closely related to their dynamic behavior. Here we use mathematical modeling to define conditions under which microtubular dynamics result in self organization. We demonstrate that random diffusional processes can generate regular microtubule organizations under specified kinetic conditions which are found to be compatible with the known properties of tubulin polymers. The organizing forces are the tubulin concentration gradients which are generated by the polymer growth. The present analysis has been restricted to the phase of establishment of the microtubule network. The same conceptual framework, applied to steady state pattern maintenance suggests that the kinetic requirements for self organization might ultimately be responsible for such extraordinary in vivo microtubule dynamics, as the rapid turnover and “dynamic instability” of the interphase network.     

A neural network model on self-organizing emergence of simple-cell receptive field with orientation selectivity in visual cortex

In order to probe into the self-organizing emergence of simple cell orientation selectivity, we tried to construct a neural network model that consists of LGN neurons and simple cells in visual cortex and obeys the Hebbian learning rule. We investigated the neural coding and representation of simple cells to a natural image by means of this model. The results show that the structures of their receptive fields are determined by the preferred orientation selectivity of simple cells. However, they are also decided by the emergence of self-organization in the unsupervision learning process. This kind of orientation selectivity results from dynamic self-organization based on the interactions between LGN and cortex.     

A neural network model on self-organizing emergence of simple-cell receptive field with orientation selectivity in visual cortex

In order to probe into the self-organizing emergence of simple cell orientation selectivity, we tried to construct a neural network model that consists of LGN neurons and simple cells in visual cortex and obeys the Hebbian learning rule. We investigated the neural coding and representation of simple cells to a natural image by means of this model. The results show that the structures of their receptive fields are determined by the preferred orientation selectivity of simple cells. However, they are also decided by the emergence of self-organization in the unsupervision learning process. This kind of orientation selectivity results from dynamic self-organization based on the interactions between LGN and cortex.