Dynamics of the ganglion cell response in the catfish and frog retinas

Responses were evoked from ganglion cells in catfish and frog retinas by a Gaussian modulation of the mean luminance. An algorithm was devised to decompose intracellularly recorded responses into the slow and spike components and to extract the time of occurrence of a spike discharge. The dynamics of both signals were analyzed in terms of a series of first-through third-order kernels obtained by cross-correlating the slow (analog) or spike (discrete or point process) signals against the white-noise input. We found that, in the catfish, (a) the slow signals were composed mostly of postsynaptic potentials, (b) their linear components reflected the dynamics found in bipolar cells or in the linear response component of type-N (sustained) amacrine cells, and (c) their nonlinear components were similar to those found in either type-N or type-C (transient) amacrine cells. A comparison of the dynamics of slow and spike signals showed that the characteristic linear and nonlinear dynamics of slow signals were encoded into a spike train, which could be recovered through the cross-correlation between the white-noise input and the spike (point process signals. In addition, well-defined spike correlates could predict the observed slow potentials. In the spike discharges from frog ganglion cells, the linear (or first-order) kernels were all inhibitory, whereas the second-order kernels had characteristics of on-off transient excitation. The transient and sustained amacrine cells similar to those found in catfish retina were the sources of the nonlinear excitation. We conclude that bipolar cells and possibly the linear part of the type-N cell response are the source of linear, either excitatory or inhibitory, components of the ganglion cell responses, whereas amacrine cells are the source of the cells' static nonlinearity.     

Interactions leading to colour opponency in ganglion cells of the turtle retina

In the turtle retina, colour-dependent photoresponses could be recorded intracellularly from ganglion cells receiving only bipolar cell input. Thus, the mechanism for colour discrimination by these ganglion cells (type A) is contained in the outer plexiform layer of the retina and depends on interaction between horizontal and cone cells. Ganglion cells receiving an additional amacrine input (type B) are not influenced by colour, and have about 0.7 logarithmic unit lower absolute sensitivity to peak wavelength than have type A ganglion cells.     

The dynamic nonlinear behavior of fly photoreceptors evoked by a wide range of light intensities.

Fly photoreceptor cells were stimulated with steps of light over a wide intensity range. First- and second-order Volterra kernels were then computed from sequences of combined step responses. Diagonal values of the second-order Volterra kernels were much greater than the off-diagonal values, and the diagonal values were roughly proportional to the corresponding first-order kernels, suggesting that the response could be approximated by a static nonlinearity followed by a dynamic linear component (Hammerstein model). The amplitudes of the second-order kernels were much smaller in light-adapted than in dark-adapted photoreceptors. Hammerstein models constructed from the step input/output measurements gave reasonable approximations to the actual photoreceptor responses, with light-adapted responses being relatively better fitted. However, Hammerstein models could not account for several features of the photoreceptor behavior, including the dependence of the step response shape on step amplitude. A model containing an additional static nonlinearity after the dynamic linear component gave significantly better fits to the data. These results indicate that blowfly photoreceptors have a strong early gain control nonlinearity acting before the processes that create the characteristic time course of the response, in addition to the nonlinearities caused by membrane conductances.     

White noise analysis of a chromatic type horizontal cell in the Xenopus retina

The dynamics of color-coded signal transmission in the light-adapted Xenopus retina were studied by a combination of white noise (Wiener) analysis and simultaneous recordings from two types of horizontal cells: chromatic-type horizontal cells (C-HCs) are hyperpolarized by blue light and depolarized by red light, whereas luminosity-type horizontal cells (L-HCs) are hyperpolarized by all wave-lengths. The retina was stimulated by two superimposed fields of red and blue light modulated by two independent white noise signals, and the resulting intracellular responses were decomposed into red and blue components (first-order kernels). The first-order kernels predict the intracellular responses with a small degree of error (3.5-9.5% in terms of mean square error) under conditions where modulated responses exceeded 30 mV in amplitude peak-to-peak, thus demonstrating that both red and blue modulation responses are linear. Moreover, there is little or no interaction between the red- and blue-evoked responses; i.e., nearly identical first-order kernels were obtained for one color whether the other color was modulated or not. In C-HCs (but not L-HCs), there were consistent differences in the dynamics of the red and blue responses. In the C-HC, the cutoff frequency of the red response was higher than for the blue (approximately 12 vs 5 Hz), and the red kernel was more bandpass than the blue. In the L-HC, kernel waveform and cutoff frequencies were similar for both colors (approximately 12 Hz or greater), and the time-to-peak of the L-HC kernel was always shorter than either the red or blue C-HC kernel. These results have implications for the mechanisms underlying color coding in the distal retina, and they further suggest that nonlinear phenomena, such as voltage-dependent conductances in HCs, do not contribute to the generation of modulation responses under the experimental conditions used here.     

Spike discharges of skeletomotor neurons during random noise modulated transmembrane current stimulation and muscle stretch

 Spike discharges of skeletomotor neurons innervating triceps surae muscles elicited by white noise modulated transmembrane current stimulation and muscle stretch were studied in decerebrated cats. The white noise modulated current intensity ranged from 4.3 to 63.2 nA peak-to-peak, while muscle stretches ranged from 100 μm to 4.26 mm peak-to-peak. The neuronal responses were studied by averaging the muscle length records centered at the skeletomotor action potentials (peri-spike average, PSA) and by Wiener analysis. Skeletomotor spikes appeared after a sharp peak in PSA of the injected current, preceded by a longer-lasting smaller wavelet of either depolarizing or hyperpolarizing direction. The PSA amplitude was not related to the injected current amplitude nor showed any differences related to the motor unit type. The PSA amplitudes were virtually independent of the stretching amplitude σ, after an initial increase with stretching amplitudes in the range of 15–40 μm (S.D.), or 100–270 μm peak-to-peak.Analyses of cross-spectra indicated a small or absent increase in gain with frequency in response to injected current, but about 20 dB/decade in the range 10–100 Hz in response to muscle stretch. The peaks of both Wiener kernels in response to current injection appear to decrease with the amplitude of injected current, but this decrease was not statistically significant. The narrow first-order kernels suggest that the transfer function between the current input and spike discharge is lowpass with a wide passband, i.e. there is very little change in dynamics. The values of the second-order kernels appear to be nonzero only along the main diagonal. This is characteristic of a simple Hammerstein type cascade, i.e. a zero memory nonlinearity followed by a linear system. Small values of second-order kernels away from the origin and narrow first-order kernels suggest that the linear cascade contributes very little to the overall dynamic response.In contrast to Wiener kernels found in response to current injection, the Wiener kernels in response to stretch showed a decreasing trend with stretch amplitude. The size of the second-order kernels decreased to a somewhat larger extent with input amplitude than that of the first-order kernels, indicating an amplitude-dependent nonlinearity. Overall, the transformation between length and spike output was described as an LNNL cascade with second-order nonlinearities. Received: 1 April 1993/Accepted in revised form: 24 March 1994     

Dynamics of turtle horizontal cell response

The small- and large-field (cone) horizontal cells produce similar dynamic responses to a stimulus whose mean luminance is modulated by a white-noise signal. Nonlinear components increase with an increase in the mean luminance and may produce a mean square error (MSE) of up to 15%. Increases in the mean luminance of the field stimulus bring about three major changes: the incremental sensitivity defined by the amplitude of the kernels decreases in a Weber-Fechner fashion; the waveforms of the kernels are transformed from monophasic (integrating) to biphasic (differentiating); the peak response time of the kernels becomes shorter and the cells respond to much higher-frequency inputs. The dynamics of the horizontal cell response also depend on the area of the retina stimulated. Smaller spots of light produce monophasic kernels of a longer peak response time. The presence of a steady background produces three major changes in the spot kernels: the kernel's amplitude becomes larger (incremental sensitivity increases); the peak response times become shorter; the waveform of the kernels changes in a fashion similar to that observed with an increase in the mean luminance of the field stimulus. A similar enhancement in the incremental sensitivity by a steady background has also been observed in catfish, which shows that this phenomenon is a common feature of the horizontal cells in the lower vertebrate retina.     

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.     

Overlapping and specific patterns of GDNF,c-ret and GFR alpha mRNA expression in the developing chicken retina

GDNF and the GDNF receptors, c-Ret, GFR alpha 1 and 2 mRNA is expressed in the developing chicken retina. GDNF labelling was mainly found in embryonic day 4-5 retina but weak labelling could also be found over scattered retinal cells at later stages. c-ret labelling was found over ganglion cells, amacrine and horizontal cells; the preferred GDNF receptor (GFR alpha 1) over amacrine and horizontal cells; and the less preferred GDNF receptor (GFR alpha 2) over ganglion cells, amacrine cells and photoreceptors.     

Dynamics of cockroach ocellar neurons

The incremental responses from the second-order neurons of the ocellus of the cockroach, Periplaneta americana, have been measured. The stimulus was a white-noise-modulated light with various mean illuminances. The kernels, obtained by cross-correlating the white-noise input against the resulting response, provided a measure of incremental sensitivity as well as of response dynamics. We found that the incremental sensitivity of the second-order neurons was an exact Weber-Fechner function; white-noise-evoked responses from second-order neurons were linear; the dynamics of second-order neurons remain unchanged over a mean illuminance range of 4 log units; the small nonlinearity in the response of the second-order neuron was a simple amplitude compression; and the correlation between the white-noise input and spike discharges of the second-order neurons produced a first-order kernel similar to that of the cell's slow potential. We conclude that signal processing in the cockroach ocellus is simple but different from that in other visual systems, including vertebrate retinas and insect compound eyes, in which the system's dynamics depend on the mean illuminance.     

牛蛙视网膜诱导型一氧化氮合酶免疫组化定位

用免疫组织化学方法研究了诱导型一氧化氮酶（iNOS)在牛蛙视网膜中的表达。结果显示,在正常状态视网膜中,无长突细胞呈弱阳性反应;节细胞层、双极细胞,水平细胞和光感受器内段呈阴性反应,在暗适应状态下,神经节细胞,内核层的无长突细胞呈强阳性反应;一些双极细胞,水平细胞和光感受器内段呈弱阳性反应,提示NO主要在暗适应状态下参与视网膜的信息传递过程。     

The nonlinear pathway of Y ganglion cells in the cat retina

Retinal ganglion cells of the Y type in the cat retina produce two different types of response: linear and nonlinear. The nonlinear responses are generated by a separate and independent nonlinear pathway. The functional connectivity in this pathway is analyzed here by comparing the observed second-order frequency responses of Y cells with predictions of a "sandwich model" in which a static nonlinear stage is sandwiched between two linear filters. The model agrees well with the qualitative and quantitative features of the second-order responses. The prefilter in the model may well be the bipolar cells and the nonlinearity and postfilter in the model are probably associated with amacrine cells.     

N-Methyl-d-aspartate receptors in the retina

The vertebrate retina is a “genuine neural center” (Ramón y Cajal), in which glutamate is a major excitatory neurotransmitter. Both N-methyl-d-aspartate (NMDA) and non-NMDA receptors are expressed in the retina. Although non-NMDA receptors and/or metabotropic glutamate receptors are generally thought to be responsible for mediating the transfer of visual signals in the outer retina, there is recent evidence suggesting that NMDA receptors are also expressed in photoreceptors, as well as horizontal and bipolar cells. In the inner retina, NMDA receptors, in addition to other glutamate receptor subtypes, are abundantly expressed to mediate visual signal transmission from bipolar cells to amacrine and ganglion cells, and could be involved in modulation of inhibitory feedback from amacrine cells to bipolar cells. NMDA receptors are extrasynaptically expressed in ganglion cells (and probably amacrine cells) and may play physiological roles in a special mode. Activity of NMDA receptors may be modulated by neuromodulators, such as d-serine and others. This article discusses retinal excitotoxicity mediated by NMDA receptors.     

Transformation of Stimulus Correlations by the Retina

Redundancies and correlations in the responses of sensory neurons may seem to waste neural resources, but they can also carry cues about structured stimuli and may help the brain to correct for response errors. To investigate the effect of stimulus structure on redundancy in retina, we measured simultaneous responses from populations of retinal ganglion cells presented with natural and artificial stimuli that varied greatly in correlation structure; these stimuli and recordings are publicly available online. Responding to spatio-temporally structured stimuli such as natural movies, pairs of ganglion cells were modestly more correlated than in response to white noise checkerboards, but they were much less correlated than predicted by a non-adapting functional model of retinal response. Meanwhile, responding to stimuli with purely spatial correlations, pairs of ganglion cells showed increased correlations consistent with a static, non-adapting receptive field and nonlinearity. We found that in response to spatio-temporally correlated stimuli, ganglion cells had faster temporal kernels and tended to have stronger surrounds. These properties of individual cells, along with gain changes that opposed changes in effective contrast at the ganglion cell input, largely explained the pattern of pairwise correlations across stimuli where receptive field measurements were possible.     

Response dynamics and receptive-field organization of catfish ganglion cells [published erratum appears in J Gen Physiol 1995 Aug;106(2):following 388]

Responses from catfish retinal ganglion cells were evoked by a spot or an annulus of light and were analyzed by a procedure identical to the one used previously to study catfish amacrine cells (Sakai H. M., and K.-I. Naka, 1992. Journal of Neurophysiology. 67:430-442.). In two- input white-noise experiments, a response evoked by simultaneous stimulation of the center and surround was decomposed into the components generated by the center and surround through a process of cross-correlation. The center and surround responses were also decomposed into their linear and nonlinear components so that the response dynamics of the linear and nonlinear components could be measured. We found that the concentric organization of the receptive field was determined by linear components, i.e., the first-order kernels generated by the center and surround were of opposite polarity. Both the center and surround generated second-order kernels with similar signatures, i.e., the second-order components formed a monotonic receptive field. The peak response time of the first- and second-order kernels from the surround was longer by approximately 20 ms than that of the center. Except for the DC potential present in the intracellular responses, almost identical first- and second-order kernels for the center and surround were obtained from both the intracellular response and spike discharges. Thus, information on concentric organization of a receptive field is translated into spike discharges with little loss of information. A train of spike discharges carries, simultaneously, at least four kinds of information: two linear and two nonlinear components, which originate in the receptive field center and the surround. A spike train is not a simple signaling device but is a carrier of complex and multiple signals. Victor, J. D., and R. M. Shapley (1979. Journal of General Physiology. 74:671-687.) discovered similarly that, in the cat retina, static second-order nonlinearity is encoded into spike trains. Results obtained in this study support the thesis that signals generated by the preganglionic cells are translated into spike discharges without major modification and that those signals can be recovered from the spike trains (Sakuranaga, M., Y. Ando, and K.-I. Naka. 1987. Journal of General Physiology. 90:229-259.; Korenberg, M. J., H. M. Sakai, and K.-I. Naka. 1989. Journal of Neurophysiology. 61:1110-1120.). Current injection studies have shown that such signal transmission is possible (Sakai, H. M., and K.-I. Naka, 1988a. Journal of Neurophysiology. 60:1549-1567.; 1990. Journal of Neurophysiology. 63:105-119.).     

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.     

Fast and slow contrast adaptation in retinal circuitry

The visual system adapts to the magnitude of intensity fluctuations, and this process begins in the retina. Following the switch from a low-contrast environment to one of high contrast, ganglion cell sensitivity declines in two distinct phases: a fast change occurs in <0.1 s, and a slow decrease over approximately 10 s. To examine where these modulations arise, we recorded intracellularly from every major cell type in the salamander retina. Certain bipolar and amacrine cells, and all ganglion cells, adapted to contrast. Generally, these neurons showed both fast and slow adaptation. Fast effects of a contrast increase included accelerated kinetics, decreased sensitivity, and a depolarization of the baseline membrane potential. Slow adaptation did not affect kinetics, but produced a gradual hyperpolarization. This hyperpolarization can account for slow adaptation in the spiking output of ganglion cells.     

A spatiotemporal white noise analysis of photoreceptor responses to UV and green light in the dragonfly median ocellus

Adult dragonflies augment their compound eyes with three simple eyes known as the dorsal ocelli. While the ocellar system is known to mediate stabilizing head reflexes during flight, the ability of the ocellar retina to dynamically resolve the environment is unknown. For the first time, we directly measured the angular sensitivities of the photoreceptors of the dragonfly median (middle) ocellus. We performed a second-order Wiener Kernel analysis of intracellular recordings of light-adapted photoreceptors. These were stimulated with one-dimensional horizontal or vertical patterns of concurrent UV and green light with different contrast levels and at different ambient temperatures. The photoreceptors were found to have anisotropic receptive fields with vertical and horizontal acceptance angles of 15 degrees and 28 degrees, respectively. The first-order (linear) temporal kernels contained significant undershoots whose amplitudes are invariant under changes in the contrast of the stimulus but significantly reduced at higher temperatures. The second-order kernels showed evidence of two distinct nonlinear components: a fast acting self-facilitation, which is dominant in the UV, followed by delayed self- and cross-inhibition of UV and green light responses. No facilitatory interactions between the UV and green light were found, indicating that facilitation of the green and UV responses occurs in isolated compartments. Inhibition between UV and green stimuli was present, indicating that inhibition occurs at a common point in the UV and green response pathways. We present a nonlinear cascade model (NLN) with initial stages consisting of separate UV and green pathways. Each pathway contains a fast facilitating nonlinearity coupled to a linear response. The linear response is described by an extended log-normal model, accounting for the phasic component. The final nonlinearity is composed of self-inhibition in the UV and green pathways and inhibition between these pathways. The model can largely predict the response of the photoreceptors to UV and green light.     

GABA release induced by aspartate-mediated activation of NMDA receptors is modulated by dopamine in a selective subpopulation of amacrine cells

Glutamate and GABA are the major excitatory and inhibitory neurotransmitters in the CNS, including the retina. In the chick retina, GABA is located in horizontal and amacrine cells and in some cells in the ganglion cell layer. It has been shown that glutamate and its agonists, NMDA, kainate, and aspartate, promote the release of GABA from isolated retina and from cultured retinal cells. Dopamine, the major catecholamine in the retina, inhibits the induction of GABA release by NMDA. Two to seven-day-old intact chicken retinas were stimulated with different glutamatergic agonists and the GABA remaining in the tissue was detected by immunohistochemical procedures. The exposure of retinas to 100 μ M NMDA for 30 minutes resulted in 50% reduction in the number of GABA-immunoreactive amacrine cells. Aspartate (100 μ M) treatment also resulted in 60% decrease in the number of GABA-immunoreactive amacrine cells. The number of GABA-immunoreactive horizontal cells was not affected by either NMDA or aspartate. In addition, dopamine reversed by 50% the reduction of the number of GABA-immunoreactive amacrine cells exposed to NMDA or aspartate. Kainate stimulation promoted a 50% reduction in the number of both GABA-immunoreactive amacrine and horizontal cells. Dopamine did not interfere with the kainate effect. While in control and in non-stimulated retinas a continuous and homogeneous immunolabeling was observed throughout the inner plexiform layer, retinas exposed to NMDA, kainate and aspartate displayed only a faint punctate labeling in the inner plexiform layer. It is concluded that, under our experimental conditions, both NMDA and aspartate induce the release of GABA exclusively from amacrine cells, and that the release is modulated by dopamine. On the other hand, kainate stimulates GABA release from both amacrine and horizontal cells with no interference of dopamine.