Spatial properties of goldfish ganglion cells

We systematically classified goldfish ganglion cells according to their spatial summation properties using the same techniques and criteria used in cat and monkey research. Results show that goldfish ganglion cells can be classified as X-, Y-, or W-like based on their responses to contrast-reversal gratings. Like cat X cells, goldfish X-like cells display linear spatial summation. Goldfish Y-like cells, like cat Y cells, respond with frequency doubling at all spatial positions when the contrast-reversal grating consists of high spatial frequencies. There is also a third class of neurons, which is neither X- nor Y-like; many of these cells' properties are similar to those of the "not-X" cells found in the eel retina. Spatial filtering characteristics were obtained for each cell by drifting sinusoidal gratings of various spatial frequencies and contrasts across the receptive field of the cell at a constant temporal rate. The spatial tuning curves of the cell depend on the temporal parameters of the stimulus; at high drift rates, the tuning curves lose their low spatial frequency attenuation. To explore this phenomenon, temporal contrast response functions were derived from the cells' responses to a spatially uniform field whose luminance varied sinusoidally in time. These functions were obtained for the center, the surround, and the entire receptive field. The results suggest that differences in the cells' spatial filtering across stimulus drift rate are due to changes in the interaction of the center and surround mechanisms; at low temporal frequencies, the center and surround responses are out-of-phase and mutually antagonistic, but at higher temporal rates their responses are in-phase and their interaction actually enhances the cell's responsiveness.     

Synaptic inputs to the ganglion cells in the tiger salamander retina

The postsynaptic potentials (PSPs) that form the ganglion cell light response were isolated by polarizing the cell membrane with extrinsic currents while stimulating at either the center or surround of the cell's receptive field. The time-course and receptive field properties of the PSPs were correlated with those of the bipolar and amacrine cells. The tiger salamander retina contains four main types of ganglion cell: "on" center, "off" center, "on-off", and a "hybrid" cell that responds transiently to center, but sustainedly, to surround illumination. The results lead to these inferences. The on-ganglion cell receives excitatory synpatic input from the on bipolars and that synapse is "silent" in the dark. The off-ganglion cell receives excitatory synaptic input from the off bipolars with this synapse tonically active in the dark. The on-off and hybrid ganglion cells receive a transient excitatory input with narrow receptive field, not simply correlated with the activity of any presynaptic cell. All cell types receive a broad field transient inhibitory input, which apparently originates in the transient amacrine cells. Thus, most, but not all, ganglion cell responses can be explained in terms of synaptic inputs from bipolar and amacrine cells, integrated at the ganglion cell membrane.     

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.     

Temporal pattern discrimination within the receptive field of cat retinal ganglion cells

ON-center and OFF-center receptive fields of cat retinal ganglion cells can be divided into two categories: sensitive (type N) and insensitive (type L) to three statistical temporal visual stimuli with different second order statistics but identical first order statistics (Tsukada et al. 1982). The temporal pattern sensitivity of type N response is closely related to the nonlinear stage of Y cells depending on the interaction between center and surround mechanism. The temporal pattern sensitivity of type N responses has a spatial profile within the receptive field; it is highly sensitive in the center region of the receptive field and less sensitive toward the field periphery. The temporal pattern sensitivity in the center region of the receptive field to statistical properties (irregular or regular) of a surrounding flash annulus shows modulation like a switching element: when the surrounding area is stimulated by a more regular flash stimulus with normal distribution of inter-stimulus intervals the system is sensitive (switching on) to the temporal pattern, while a change to an irregular one with an exponential distribution makes it insensitive (switching off) to the temporal pattern.     

Spatial asymmetries in cat retinal ganglion cell responses

. Enroth-Cugell and Robson (1966) first proposed a classification of retinal ganglion cells into X cells, which exhibit approximate linear spatial summation and largely sustained responses, and Y cells, which exhibit nonlinearities and transient responses. Gaudiano (1992a, 1992b, 1994) has suggested that the dominant characteristics of both X and Y cells can be simulated with a single model simply by changing receptive field profiles to match those of the anatomical counterparts of X and Y cells. He also proposed that a significant component of the spatial nonlinearities observed in Y (and sometimes X) cells can result from photoreceptor nonlinearities coupled with push-pull bipolar connections. Specifically, an asymmetry was predicted in the ganglion cell response to rectangular gratings presented at different locations in the receptive field under two conditions: introduction/withdrawal (on-off) or contrast reversal. When measuring the response to these patterns as a function of spatial phase, the standard difference-of-Gaussians model predicts symmetrical responses about the receptive field center, while the push-pull model predicts slight but significant asymmetry in the on-off case only. To test this hypothesis, we have recorded ganglion cell responses from the optic tract fibers of anesthetized cat. The mean and standard deviations of responses to on-off and contrast-reversed patterns were compared. We found that all but one of the cells that yielded statistically significant data confirmed the hypothesis. These results largely support the theoretical prediction. Received: 21 March 1997 / Accepted in revised form: 6 May 1998     

The effect of strychnine, bicuculline, and picrotoxin on X and Y cells in the cat retina

The effect of intravenous strychnine and the GABA antagonists picrotoxin and bicuculline upon the discharge pattern of center-surround-organized cat retinal ganglion cells of X and Y type were studied. Stimuli (mostly scotopic, and some photopic) were selected such that responses from both on and off-center cells were either due to the center, due to the surround, or clearly mixed. Pre-drug control responses were obtained, and their behavior following administration of the antagonists was observed for periods up to several hours. X-cell responses were affected in a consistent manner by strychnine while being unaffected by GABA antagonists. All observed changes following strychnine were consistent with a shift in center-surround balance of X cells in favor of the center. For Y-cell responses to flashing annuli following strychnine, there was either no shift or a relatively small shift in center-surround balance. Compared to X-cell responses to flashing lights, those of Y cells were very little affected by strychnine and in most cases were unaffected. It thus appears that glycine plays a similar role in receptive field organization of X cells as does GABA in Y cells (Kirby and Enroth-Cugell, 1976. J. Gen. Physiol. 68:465-484).     

Receptive field mechanisms of ganglion cells in the cat retina

On the basis of anatomical and physiological results of the vertebrate retina, a method is proposed for analysing the respective fields of ganglion cells in the cat retina. In the model, we assume the following: (a) Ganglion cells receive their input from bipolar and/or amacrine cells. (b) The nonlinearity of ganglion cell responses is due to the activities of transient type amacrine cells. The method has been proved to be effective. According to the results of this investigation, the receptive field properties of X type and Y type ganglion cells are heterogeneous. Thus, it may be considered that their receptive fields consist of center and surround mechanisms. The receptive field properties of X-cells are almost linear and the X-cells seem to receive most of their input from bipolar cells. On the other hand, the ones of Y-cells are highly nonlinear. Consequently, it is conceivable that the Y-cells receive their input mainly from transient type amacrine cells.     

Receptive field mechanisms of cat X and Y retinal ganglion cells

We investigated receptive field properties of cat retinal ganglion cells with visual stimuli which were sinusoidal spatial gratings amplitude modulated in time by a sum of sinusoids. Neural responses were analyzed into the Fourier components at the input frequencies and the components at sum and difference frequencies. The first-order frequency response of X cells had a marked spatial phase and spatial frequency dependence which could be explained in terms of linear interactions between center and surround mechanisms in the receptive field. The second-order frequency response of X cells was much smaller than the first-order frequency response at all spatial frequencies. The spatial phase and spatial frequency dependence of the first-order frequency response in Y cells in some ways resembled that of X cells. However, the Y first-order response declined to zero at a much lower spatial frequency than in X cells. Furthermore, the second-order frequency response was larger in Y cells; the second-order frequency components became the dominant part of the response for patterns of high spatial frequency. This implies that the receptive field center and surround mechanisms are physiologically quite different in Y cells from those in X cells, and that the Y cells also receive excitatory drive from an additional nonlinear receptive field mechanism.     

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.     

On the characteristic of nonlinear spatial summation observed in the lateral geniculate cells of cats

The spatial summation characteristic in the receptive fields of cat lateral geniculate cells were investigated. First, the central area of the receptive field was determined using a spot of light. Then the response of the cell were obtained using disc-shaped stimuli of various radii located in the middle point of the receptive field center. When the radius was increased gradually, the response tended to increase, at first, until it reached a peak value and began to decrease thereafter. The radius where the peak response took place was generally less than that of the receptive field center. Furthermore, this radius decreased when the intensity of the stimulus light was increased. These neurophysiological findings could be simulated by a model. The model consists of two parts. The first part receives the input from the photoreceptors. It is of homogeneous structure with shunting inhibition. The second part receives the input from the first part. The structure is characterized by the conventional center-surround type lateral interaction.     

Neural response mechanisms in the photoreceptive pineal organ of goldfish

In order to classify the different cell types involved in signal transmission of the photoreceptive pineal organ of the goldfish, Carassius auratus, intra- and extracellular electrical responses were recorded from photoreceptors and second-order neurons. Photoreceptor responses to light consisted of hyperpolarizing potentials up to 30 mV. The responses were graded with intensity and their voltage-intensity relation followed the hyperbolic function V/Vmax = In/In + sigma n. Latencies varied between 500 msec for responses near threshold and 60 msec for supersaturating flashes. The response duration increased up to 60 sec for flashes 2 log units above the saturation level. Action spectra of individual photoreceptors peaked at lambda max = 530 nm and corresponded to measurements of extracellular slow mass potentials or spike potentials. Slow mass potentials exhibited similar characteristics as intracellular recorded photoreceptor potentials with respect to latency, voltage-intensity curves and spectral sensitivity. Ganglion cells showed maintained discharges under conditions of steady illumination. The discharge rate changed inversely with the logarithm of steady illumination over a range of 8 log units. The response to light flashes was purely achromatic and consisted of inhibition of the maintained discharge. The physiological properties demonstrate that the pineal organ of the goldfish is an effective functional photoreceptor organ operating both in dim and in bright light. The light-induced hyperpolarization of photoreceptors lead to an inhibition of the nervous discharge of ganglion cells. The direct flow of information from photoreceptors to ganglion cells is the basic channel of data processing in the goldfish pineal.     

Receptive field microstructure and dendritic geometry of retinal ganglion cells

We studied the fine spatial structure of the receptive fields of retinal ganglion cells and its relationship to the dendritic geometry of these cells. Cells from which recordings had been made were microinjected with Lucifer yellow, so that responses generated at precise locations within the receptive field center could be directly compared with that cell's dendritic structure. While many cells with small receptive fields had domeshaped sensitivity profiles, the majority of large receptive fields were composed of multiple regions of high sensitivity. The density of dendritic branches at any one location did not predict the regions of high sensitivity. Instead, the interactions between a ganglion cell's dendritic tree and the local mosaic of bipolar cell axons seem to define the fine structure of the receptive field center.     

Differential targeting of optical neuromodulators to ganglion cell soma and dendrites allows dynamic control of center-surround antagonism

Retinal degenerative diseases cause photoreceptor loss and often result in remodeling and deafferentation of the inner retina. Fortunately, ganglion cell morphology appears to remain intact long after photoreceptors and distal retinal circuitry have degenerated. We have introduced the optical neuromodulators channelrhodopsin-2 (ChR2) and halorhodopsin (NpHR) differentially into the soma and dendrites of ganglion cells to recreate antagonistic center-surround receptive field interactions. We then reestablished the physiological receptive field dimensions of primate parafoveal ganglion cells by convolving Gaussian-blurred versions of the visual scene at the appropriate wavelength for each neuromodulator with the Gaussians inherent in the soma and dendrites. These Gaussian-modified ganglion cells responded with physiologically relevant antagonistic receptive field components and encoded edges with parafoveal resolution. This approach bypasses the degenerated areas of the distal retina and could provide a first step in restoring sight to individuals suffering from retinal disease.     

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.     

Features and functions of nonlinear spatial integration by retinal ganglion cells

Ganglion cells in the vertebrate retina integrate visual information over their receptive fields. They do so by pooling presynaptic excitatory inputs from typically many bipolar cells, which themselves collect inputs from several photoreceptors. In addition, inhibitory interactions mediated by horizontal cells and amacrine cells modulate the structure of the receptive field. In many models, this spatial integration is assumed to occur in a linear fashion. Yet, it has long been known that spatial integration by retinal ganglion cells also incurs nonlinear phenomena. Moreover, several recent examples have shown that nonlinear spatial integration is tightly connected to specific visual functions performed by different types of retinal ganglion cells. This work discusses these advances in understanding the role of nonlinear spatial integration and reviews recent efforts to quantitatively study the nature and mechanisms underlying spatial nonlinearities. These new insights point towards a critical role of nonlinearities within ganglion cell receptive fields for capturing responses of the cells to natural and behaviorally relevant visual stimuli. In the long run, nonlinear phenomena of spatial integration may also prove important for implementing the actual neural code of retinal neurons when designing visual prostheses for the eye.     

Melanopsin--shedding light on the elusive circadian photopigment

Circadian photoentrainment is the process by which the brain's internal clock becomes synchronized with the daily external cycle of light and dark. In mammals, this process is mediated exclusively by a novel class of retinal ganglion cells that send axonal projections to the suprachiasmatic nuclei (SCN), the region of the brain that houses the circadian pacemaker. In contrast to their counterparts that mediate image-forming vision, SCN-projecting RGCs are intrinsically sensitive to light, independent of synaptic input from rod and cone photoreceptors. The recent discovery of these photosensitive RGCs has challenged the long-standing dogma of retinal physiology that rod and cone photoreceptors are the only retinal cells that respond directly to light and has explained the perplexing finding that mice lacking rod and cone photoreceptors can still reliably entrain their circadian rhythms to light. These SCN-projecting RGCs selectively express melanopsin, a novel opsin-like protein that has been proposed as a likely candidate for the photopigment in these cells. Research in the past three years has revealed that disruption of the melanopsin gene impairs circadian photo- entrainment, as well as other nonvisual responses to light such as the pupillary light reflex. Until recently, however, there was no direct demonstration that melanopsin formed a functional photopigment capable of catalyzing G-protein activation in a light-dependent manner. Our laboratory has recently succeeded in expressing melanopsin in a heterologous tissue culture system and reconstituting a pigment with the 11-cis-retinal chromophore. In a reconstituted biochemical system, the reconstituted melanopsin was capable of activating transducin, the G-protein of rod photoreceptors, in a light-dependent manner. The absorbance spectrum of this heterologously expressed melanopsin, however, does not match that predicted by previous behavioral and electophysiological studies. Although melanopsin is clearly the leading candidate for the elusive photopigment of the circadian system, further research is needed to resolve the mystery posed by its absorbance spectrum and to fully elucidate its role in circadian photoentrainment.     

Movement sensitivities of cells in the fly's medulla

Summary Intracellular recordings were made in the medullae of intact, restrained females ofCalliphora vicina that faced a hemispherical, minimum-distortion surface upon which moving patterns and spots were projected from the rear (Fig. 2). In the distal medulla, noisy hyperpolarizations to light, most likely recorded in terminals of laminar (L) cells, had flicker-like oscillations to moving gratings of 15° spatial wavelength but not of 2.5° spatial wavelength (Fig. 3). Medullary (M) cells penetrated distally responded to grating movements with similar but depolarizing oscillations, in one cell 180° out of phase with a nearby laminar response (Figs. 4–6).A characteristic movement response recorded from most medullary cells consisted of abrupt, maintained nondirectional depolarizations in response to movements of gratings, often with directional ripple or spikes superimposed. When directions of movement reversed, there were brief repolarizations, but when movements stopped, depolarizations decayed away more slowly (Figs. 7 and 8). Magnitude of responses increased with increasing speeds of both 15° and 2.5° gratings (Figs. 9–11). In some cells, there were delayed decays of responses after stopping (Fig. 12). Still other cells seemed to receive inhibition from other, characteristically responding cells (Fig. 13).Receptive fields tested were simple and usually large, with only a suggestion of surround inhibition (Fig. 14). In general, intensity and position were interchangeable over a cell's receptive field (Figs. 15 and 16). Moving edges and dark spots elicited responses primarily within receptive field centers (Figs. 18–20).It is argued that waveforms of characteristic movement responses can be explained by multiplicative inputs from L- and M-cells to movement detectors (Figs. 21–26).Abbreviations L cells laminar (monopolar) cells - M cells medullary cells     

GABA及荷包牡丹碱对金黄地鼠上丘视觉神经...

Visual response properties of single neurons in the superior colliculus of golden hamsters could be altered by iontophoretically applied gamma-aminobutyric acid (GABA) and its antagonist, bicuculline (Bicu). GABA decreased the responses of the superficial cells to stationary flashing stimuli, while Bicu increased the responses and suppressed the inhibition exerted by the surround. The number of spikes evoked by a moving bar/spot decreased after applying GABA in 76.6% of the cells (n = 60) and increased in 90.0% (n = 60) of the cells after Bicu. Similar effects on the spontaneous activities were observed. In addition, 65.0% of the 60 cells recorded have enlarged movement receptive fields after application of Bicu. GABA and Bicu have some effects on the orientation selectivity of the collicular cells too.     

Characteristics of dorsal horn neurons expressing subliminal responses to sural nerve stimulation

The present study was designed (1) to characterize the subliminal responses of dorsal horn neurons to stimulation of the sural nerve, and (2) to correlate the type of response to this stimulus with the responses to natural mechanical stimulation of the skin. To accomplish this, intracellular and extracellular recordings were carried out in L6 and L7 dorsal horn neurons in the cat. The excitatory responses of each cell to electrical stimulation of the sural nerve and to mechanical stimulation of the skin were noted. Of 35 dorsal horn cells recorded intracellularly, 11 responded with impulses to sural nerve stimulation, 9 responded with excitatory postsynaptic potentials (EPSPs) but not impulses, and 15 had no excitatory responses to this stimulus. The type of response to sural nerve stimulation was strongly correlated with receptive field modality. Most cells receiving an input from high-threshold cutaneous mechanoreceptors responded with impulses or gave no excitatory response to sural nerve stimulation, whereas most cells that had only low-threshold mechanoreceptor input responded with EPSPs only or gave no response. In cells with only low-threshold (LT) mechanoreceptive input, response to sural nerve stimulation was highly correlated with receptive field locus. Those LT cells with no excitatory responses to sural nerve stimulation had receptive fields confined to the foot and/or toes, whereas those that gave EPSPs had more proximal receptive fields. The possible significance of these data with reference to changes observed after lesions, such as increased response to sural nerve stimulation, increased receptive field size, and somatotopic reorganization, is discussed.     

用菌紫质Langmuir—Blodgett膜模拟视网膜神经节细胞感受野的DOG滤波等特性

用菌紫质ＬＢ（Ｌａｎｇｍｕｉｒ—Ｂｌｏｄｇｅｔｔ）膜以一维形式模拟了视网膜神经节细胞的ＯＮ－中心型感受野。实验表明菌紫质ＬＢ膜具有ＯＮ型和ＯＦＦ型微分响应特性。对运动狭缝,所模拟的人工视网膜感受野的周边区和中心区都具有类高斯函数形式的滤波特性,整个人工视网膜感受野具有与高等动物视网膜相似的ＤＯＧ（ＤｉｆｆｅｒｅｎｃｅｏｆＧａｕｓｓｉａｎｓ）滤波运算功能。