Processing of Color- and Noncolor-Coded Signals in the Gourami Retina. III. Ganglion Cells

1997 ◽  
Vol 78 (4) ◽  
pp. 2034-2047 ◽  
Author(s):  
Hiroko M. Sakai ◽  
Hildred Machuca ◽  
Michael J. Korenberg ◽  
Ken-Ichi Naka
Keyword(s):  
White Noise ◽  
Spike Train ◽  
Ganglion Cells ◽  
Green Light ◽  
Amacrine Cells ◽  
Second Order ◽  
First Order ◽  
Wide Range ◽  
Mean Luminance ◽  
Order Components

Sakai, Hiroko M., Hildred Machuca, Michael J. Korenberg, and Ken-Ichi Naka. Processing of color- and noncolor-coded signals in the gourami retina. III. Ganglion cells. J. Neurophysiol. 78: 2034–2047, 1997. The dynamics of intracellular responses from ganglion cells, as well as that of spike discharges, were studied with the stimulus regimens and analytic procedures identical to those used to study the dynamics of the responses from horizontal and amacrine cells ( Sakai et al. 1997a , b ). The stimuli used were large fields of red and green light given as a pulsatile input or modulation about a mean luminance by a white-noise signal. Spike discharges evoked by a white-noise stimulus were analyzed in exactly the same manner as that used for analysis of analog responses. The canonical nature of kernels allowed us to correlate the first- and second-order components in a spike train with those of the intracellular responses from horizontal, amacrine, and ganglion cells. Both red and green stimuli given alone in darkness produced noncolor-coded responses from all ganglion cells. In the case of some cells, steady red illumination changed the polarity or waveform of the response to green light. Color-coded ganglions responded only to simultaneous color contrast. Nonlinearities recovered from intracellular responses, and spike discharges were similar to those found in responses from amacrine cells and were of two types, one characteristic of the C amacrine cells and the other characteristic of the N amacrine cells. The first-order kernels of most ganglion cells could be divided into two basic types, biphasic and triphasic. The combination of kernels of these two basic types with different polarities can produce a wide range of responses. Addition of two types of second-order nonlinearity could render color coding in this relatively simple retina as an extremely complex process. Color information appeared to be represented by the polarity, as well as the waveform, of the first-order kernel. The response dynamics is a means of transmission of color-coded information. Second-order components carry information about changes around a mean luminance regardless of the color of an input. Some spike discharges produced a well-defined cross-kernel between red and green inputs to show that a particular time sequence of red and green stimuli was detected by the retinal neuron network. The similarity between signatures of second-order kernels for both amacrine and ganglion cells indicates that signals undergo a minimal transformation in the temporal domain when they are transmitted from amacrine to ganglion cells and then transformed into a spike train. Under our experimental conditions, a single spike train carried simultaneously information about red and green inputs, as well as about linear and nonlinear components. In addition, the spike train also carries a cross-talk component. A spike train is a carrier of multiple signals. Conversely, many types of information in a stimulus are independently encoded into a spike train.

Response dynamics and receptive-field organization of catfish amacrine cells

1992 ◽  
Vol 67 (2) ◽  
pp. 430-442 ◽  
Author(s):  
H. M. Sakai ◽  
K. Naka
Keyword(s):  
Receptive Field ◽  
White Noise ◽  
Amacrine Cells ◽  
Second Order ◽  
Response Dynamics ◽  
Field Center ◽  
Mean Luminance ◽  
Nonlinear Components ◽  
Receptive Field Center ◽  
Dc Potentials

1. We have applied Wiener analysis to a study of response dynamics of N (sustained) and C (transient) amacrine cells. Stimuli were a spot and an annulus of light, the luminance of which was modulated by two independent white-noise signals. First- and second-order Wiener kernels were computed for each spot and annulus input. The analysis allowed us to separate a modulation response into its linear and nonlinear components, and into responses generated by a receptive-field center and its surround. 2. Organization of the receptive field of N amacrine cells consists of both linear and nonlinear components. The receptive field of linear components was center-surround concentric and opposite in polarity, whereas that of second-order nonlinear components was monotonic. 3. In NA (center-depolarizing) amacrine cells, the membrane DC potentials brought about by the mean luminance of a white-noise spot or a steady spot were depolarizations, whereas those brought about by the mean luminance of a white-noise annulus or a steady annulus were hyperpolarizations. In NB (center-hyperpolarizing) amacrine cells, this relationship was reversed. If both receptive-field center and surround were stimulated by a spot and annulus, membrane DC potentials became close to the dark level and the amplitude of modulation responses became larger. 4. The linear responses of a receptive-field center of an N amacrine cell, measured in terms of the first-order Wiener kernel, were facilitated if the surround was stimulated simultaneously. The amplitude of the kernel became larger, and its peak response time became shorter. The same facilitation occurred in the linear responses of a receptive-field surround if the center was stimulated simultaneously. 5. The second-order nonlinear responses were not usually generated in N amacrine cells if the stimulus was either a white-noise spot or a white-noise annulus alone. Significant second-order nonlinearity appeared if the other region of the receptive field was also stimulated. 6. Membrane DC potentials of C amacrine cells remained at the dark level with either a white-noise spot or a white-noise annulus. The cell responded only to modulations. 7. The major characteristics of center and surround responses of C amacrine cells could be approximated by second-order Wiener kernels of the same structure. The receptive field of second-order nonlinear components of C amacrine cells was monotonic.(ABSTRACT TRUNCATED AT 400 WORDS)

Signal transmission in the catfish retina. V. Sensitivity and circuit

1987 ◽  
Vol 58 (6) ◽  
pp. 1329-1350 ◽  
Author(s):  
H. M. Sakai ◽  
K. Naka
Keyword(s):  
Ganglion Cells ◽  
Modulation Depth ◽  
Horizontal Cell ◽  
Amacrine Cells ◽  
Bipolar Cells ◽  
Modulation Response ◽  
First Order ◽  
Parametric Change ◽  
Type C ◽  
Mean Luminance

1. We analyzed the light-evoked responses of retinal neurons by means of a white-noise technique. Horizontal and bipolar cells produced a modulation response that was linearly related to a modulation of the mean luminance of a large field of light. The first-order kernels were capable of reproducing the cells' modulation response with a fair degree of accuracy. The amplitude as well as the waveform of the kernels changed with the change in the mean luminance. This is a parametric change and is a form of field adaptation. As the time constant of the parametric change was much longer than that of the modulation response (memory), neurons were assumed to be at a dynamic steady state at a given mean luminance. 2. With the presence of a steady annular illumination, the first-order kernel derived from stimulation with a small spot of light became faster in peak response time and larger in amplitude. For horizontal-cell somas and bipolar cells, the surround also linearized their modulation response. This surround enhancement has been seen in all the cone-driven retinal cells except the receptor and horizontal cell axon, in which a steady surround decreased the amplitude of the spot-evoked kernel but shortened the peak response time. 3. A change in the modulation depth did not affect either the amplitude or the wave-form of the first-order kernels from the horizontal and bipolar cells. In the amacrine and ganglion cells, on the other hand, the amplitude of kernels was related inversely to the depth of modulation. These cells were more sensitive to the modulation of a small modulation depth. 4. A static nonlinearity appeared when signals were transmitted to the amacrine cells. The nonlinearity was first produced in the type-C amacrine cells by a process, which could be modeled by squaring the bipolar cell response. A gamut of more complex second-order nonlinearities found in type-N amacrine cells could be modeled by a band-pass filtering of the type-C cell response. Linear components in the bipolar cells and nonlinear components in the amacrine cells are encoded into spike trains in the ganglion cells. Thus, under our simple stimulus regimen, the ganglion cells transformed the results of the preganglionic signal processing into a spike train without much modification. 5. We propose a tentative diagram of the signal flow in the cone-driven catfish retinal neurons based on this and previous studies.(ABSTRACT TRUNCATED AT 400 WORDS)

Processing of Color- and Noncolor-Coded Signals in the Gourami Retina. II. Amacrine Cells

1997 ◽  
Vol 78 (4) ◽  
pp. 2018-2033 ◽  
Author(s):  
Hiroko M. Sakai ◽  
Hildred Machuca ◽  
Ken-Ichi Naka
Keyword(s):  
White Noise ◽  
Amacrine Cell ◽  
Amacrine Cells ◽  
Second Order ◽  
Order Component ◽  
Response Dynamics ◽  
Dendritic Field ◽  
State Response ◽  
Order Components ◽  
Nonlinear Components

Sakai, Hiroko M., Hildred Machuca, and Ken-Ichi Naka. Processing of color- and noncolor-coded signals in the gourami retina. II. Amacrine cells. J. Neurophysiol. 78: 2018–2033, 1997. The same set of stimuli and analytic methods that was used to study the dynamics of horizontal cells ( Sakai et al. 1997a ) was applied to a study of the response dynamics and signal processing in amacrine cells in the retina of the kissing gourami, Helostoma rudolfi. The retina contains two major classes of amacrine cells that could be identified from their morphology: C and N amacrine cells. C amacrine cells had a two-layered dendritic field, whereas N cells had a monolayered dendritic field. Both types of amacrine cell were tracer-coupled but coupling was more extensive in the N amacrine cells. Responses from C amacrine cells lacked a DC component and had a small linear component that was <10% in terms of mean square error (MSE); the second-order component often accounted for >50% of the modulation response. The C amacrine cells did not show any characteristic color coding under any stimulus condition. Most responses of N cells to a pulsatile stimulus consisted of a series of depolarizing transient potentials and steady illumination did not generate any DC potential in these cells. The response to a white-noise modulated input was composed of well-defined first- and second-order components and, possibly, higher-order components. The response evoked by a red or green white-noise–modulated stimulus given alone was not color coded. Modulated red illumination in the presence of a green illumination elicited a color-coded response from >70% of N amacrine cells. Color information was carried not only by the polarity but also by the dynamics of the first-order component. No convincing evidence was obtained to indicate that the second-order component might be involved in color processing. Some N amacrine cells produced a well-defined (second-order) interaction kernel to show that the temporal sequence of red and green stimuli was a parameter to be considered. In a complex cell such as an amacrine cell, responses evoked by a pulsatile stimulus given in darkness and by modulation of a mean luminance could be very different in terms of their characteristics. It was not always possible to predict the response evoked by one stimulus from observing the cell's response to another stimulus. This is because, in N cells, a flash-evoked (nonsteady state) response is composed largely of nonlinear components whereas a modulation (steady state) response is composed of linear as well as nonlinear components.

Dissection of the neuron network in the catfish inner retina. V. Interactions between NA and NB amacrine cells

1990 ◽  
Vol 63 (1) ◽  
pp. 120-130 ◽  
Author(s):  
H. M. Sakai ◽  
K. I. Naka
Keyword(s):  
White Noise ◽  
Small Amplitude ◽  
Amacrine Cell ◽  
Amacrine Cells ◽  
The Other ◽  
First Order ◽  
Membrane Noise ◽  
Current Pulses ◽  
Damped Oscillation ◽  
Order Kernel

1. Simultaneous intracellular recordings were made from two neighboring N amacrine cells, one an ON amacrine (NA) cell and the other an OFF amacrine (NB) cell. Extrinsic current was injected into one amacrine cell, and the resulting intracellular responses were recorded from the other amacrine cell. Test signals included 1) a single-frequency sinusoid, 2) a depolarizing or hyperpolarizing pulse, or 3) a white-noise modulated current. In some cell pairs, membrane noise was measured in the dark as well as under a steady background illumination. 2. Current pulses injected into a NA cell evoked a damped oscillation from a NB cell. The first-order kernel derived by cross-correlating the white-noise current injected into a NA cell against the evoked response from a NB cell was a large depolarization followed by a damped oscillation. The frequency of oscillations varied slightly from pair to pair but averaged 35 Hz. 3. Current pulses injected into a NB cell evoked a sign-inverting response (hyperpolarization) of very small amplitude from a NA cell. Similarly, the first-order kernel was a hyperpolarization of very small amplitude. 4. The power spectrum of the membrane noise recorded from NA and NB cells in the dark or during steady illumination often showed a peak at 35 Hz. Such membrane noise synchronizes synergistically among NA cells and among NB cells in the dark. In addition, the membrane fluctuations seen in NA and NB cells in the dark were out of phase. 5. Transmission between NA and NB cells was largely accounted for by a linear component; however, a very small but significant second- and third-order nonlinearity was also generated. 6. These results show that the interactions occurring between amacrine cells of opposite response polarity are much more complex than those between cells of the same response polarity and that the neural circuitry in the inner retina actively controls interactions between ON and OFF channels in the dark as well as in the presence of light stimuli.

An Energy Method for Certain Second-Order Effects With Application to Torsion of Elastic Bars Under Tension

1980 ◽  
Vol 47 (1) ◽  
pp. 75-81 ◽  
Author(s):  
R. T. Shield
Keyword(s):  
Axial Force ◽  
Strain Energy Function ◽  
Incompressible Material ◽  
Elastic Cylinder ◽  
Second Order ◽  
The Other ◽  
Circular Cylinders ◽  
Order Effects ◽  
First Order ◽  
Wide Range

When a mechanical system has a potential energy, it is a simple matter to show that if the generalized force corresponding to a coordinate p is known to first order in p for a range of the other coordinates of the system, then the other generalized forces can be found immediately to second order in p, without requiring a second-order analysis of the system. By this method the second-order change in the axial force when a finitely extended elastic cylinder is twisted is found from the first-order value of the twisting moment. Numerical results for a realistic form of the strain-energy function for an incompressible material suggest that the second-order expression for the axial force is very accurate for a wide range of twist for circular cylinders of rubber-like materials extended 100 percent or more.

scholarly journals Dynamics of the ganglion cell response in the catfish and frog retinas.

1987 ◽  
Vol 90 (2) ◽  
pp. 229-259 ◽  
Author(s):  
M Sakuranaga ◽  
Y Ando ◽  
K Naka
Keyword(s):  
White Noise ◽  
Ganglion Cell ◽  
Point Process ◽  
Cell Response ◽  
Ganglion Cells ◽  
Linear Part ◽  
Amacrine Cells ◽  
Bipolar Cells ◽  
Response Component ◽  
Noise Input

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.

Pharmacology of Directionally Selective Ganglion Cells in the Rabbit Retina

1997 ◽  
Vol 77 (2) ◽  
pp. 675-689 ◽  
Author(s):  
Christopher A. Kittila ◽  
Stephen C. Massey
Keyword(s):  
Ganglion Cells ◽  
Amacrine Cells ◽  
Gaba Release ◽  
Excitatory Input ◽  
Directional Selectivity ◽  
Dependent Manner ◽  
Rabbit Retina ◽  
Response Curves ◽  
Wide Range ◽  
Starburst Amacrine Cells

Kittila, Christopher A. and Stephen C. Massey. Pharmacology of directionally selective ganglion cells in the rabbit retina. J. Neurophysiol. 77: 675–689, 1997. In this report we describe extracellular recordings made from on and on-off directionally selective (DS) ganglion cells in the rabbit retina during perfusion with agonists and antagonists to acetylcholine (ACh), glutamate, and γ-aminobutyric acid (GABA). Nicotinic ACh agonists strongly excited DS ganglion cell in a dose-dependent manner. Dose-response curves showed a wide range of potencies, with (±)-exo-2-(6-chloro-3pyridinyl)-7-azabicyclo[2.2.1] heptane dihydrochloride (epibatidine) ≫ nicotine > 1,1-dimethyl-4-phenylpiperazinium iodide = carbachol. In addition, the mixed cholinergic agonist carbachol produced a small excitation, mediated by muscarinic receptors, that could be blocked by atropine. The specific nicotinic antagonists hexamethonium bromide (100 μM), dihydro-β-erythroidine (50 μM), mecamylamine (50 μM), and tubocurarine (50 μM) blocked the responses to nicotinic agonists. In addition, nicotinic antagonists reduced the light-driven input to DS ganglion cells by ∼50%. However, attenuated responses were still DS. We deduce that cholinergic input is not required for directional selectivity. These experiments reveal the importance of bipolar cell input mediated by glutamate. N-methyl-d-aspartic acid (NMDA) excited DS ganglion cells, but NMDA antagonists did not abolish directional selectivity. However, a combined cholinergic and NMDA blockade reduced the responses of DS ganglion cells by >90%. This indicates that most of the noncholinergic excitatory input appears to be mediated by NMDA receptors, with a small residual made upb y  α - a m i n o - 3 - h y d r o x y - 5 - m e t h y l - 4 - i s o x a z o l e p r o p i o n i c  a c i d(AMPA)/kainate (KA) receptors. Responses to AMPA and KA were highly variable and often evoked a mixture of excitation and inhibition due to the release of ACh and GABA. Under cholinergic blockade AMPA/KA elicited a strong GABA-mediated inhibition in DS ganglion cells. AMPA/KA antagonists, such as 2,3-dihydroxy-6-nitro-7-sulfamoylbenzo(F)quinoxaline dione and GYKI-53655, promoted null responses and abolished directional selectivity due to the blockade of GABA release. We conclude that GABA release, mediated by non-NMDA glutamate receptors, is an essential part of the mechanism of directional selectivity. The source of the GABA is unknown, but may arise from starburst amacrine cells.

scholarly journals Nonlinear temporal receptive fields of neurons in the dorsal cochlear nucleus

2013 ◽  
Vol 110 (10) ◽  
pp. 2414-2425 ◽  
Author(s):  
Sharba Bandyopadhyay ◽  
Eric D. Young
Keyword(s):  
Cochlear Nucleus ◽  
Receptive Fields ◽  
Sound Level ◽  
Second Order ◽  
Dorsal Cochlear Nucleus ◽  
Spectral Processing ◽  
Low Contrast ◽  
Response Properties ◽  
First Order ◽  
Order Components

Studies of the dorsal cochlear nucleus (DCN) have focused on spectral processing because of the complex spectral receptive fields of the DCN. However, temporal fluctuations in natural signals convey important information, including information about moving sound sources or movements of the external ear in animals like cats. Here, we investigate the temporal filtering properties of DCN principal neurons through the use of temporal weighting functions that allow flexible analysis of nonlinearities and time variation in temporal response properties. First-order temporal receptive fields derived from the neurons are sufficient to characterize their response properties to low-contrast (3-dB standard deviation) stimuli. Larger contrasts require the second-order terms. Allowing temporal variation of the parameters of the first-order model or adding a component representing refractoriness improves predictions by the model by relatively small amounts. The importance of second-order components of the model is shown through simulations of nonlinear envelope synchronization behavior across sound level. The temporal model can be combined with a spectral model to predict tuning to the speed and direction of moving sounds.

scholarly journals Receptive field mechanisms of cat X and Y retinal ganglion cells.

1979 ◽  
Vol 74 (2) ◽  
pp. 275-298 ◽  
Author(s):  
J D Victor ◽  
R M Shapley
Keyword(s):  
Receptive Field ◽  
Spatial Frequency ◽  
Frequency Response ◽  
Retinal Ganglion Cells ◽  
Ganglion Cells ◽  
Second Order ◽  
Retinal Ganglion ◽  
First Order ◽  
X Cells ◽  
Y 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.

UV responses in the retina of the turtle

1999 ◽  
Vol 16 (2) ◽  
pp. 191-204 ◽  
Author(s):  
D.F. VENTURA ◽  
J.M. de SOUZA ◽  
R.D. DEVOE ◽  
Y. ZANA
Keyword(s):  
Ganglion Cells ◽  
Uv Light ◽  
Green Light ◽  
Amacrine Cells ◽  
Horizontal Cells ◽  
Stray Light ◽  
Wide Field ◽  
Beta Band ◽  
Sensitivity Curves ◽  
Response Method

To study processing of UV stimuli in the retina of the turtle, Trachemys dorbignii, we recorded intracellular responses to spectral light from 89 cells: 54 horizontal (47 monophasic, five (R/G) biphasic and two (Y/B) triphasic), 14 bipolar, 12 amacrine, and nine ganglion cells. Spectral sensitivities were measured with monochromatic flashes or with the dynamic constant response method in dark or chromatic adapted states. Stray light and second-order harmonics were also measured. (1) All cells responded to UV stimuli, although none had maximum sensitivity in the UV. (2) Most horizontal, bipolar, and amacrine cells had red-peaked spectral sensitivities. (3) Red adaptation of all monophasic horizontal cells indicated a single red input, except one that had additional peaks in the blue and UV. (4) Responses of biphasic and triphasic horizontal cells to UV light were always hyperpolarizing. Opposition between hyperpolarizing and depolarizing responses at long wavelengths indicates that UV responses were not due to the beta band of red receptors. (5) An unstained spectrally opponent bipolar cell hyperpolarized in the center to green light and antagonistically depolarized in the surround to UV, blue, and green flashes, but hyperpolarized to red. (6) All dark-adapted amacrine cells were red-peaked monophasic cells, but red adaptation broadened their spectral-sensitivity curves or displaced their peaks. An A15, an A18, and an A24 wide-field amacrine cell were stained. (7) A G15 bistratified ganglion cell is shown here for the first time to be spectrally opponent. This UVB/RG cell depolarized to UV and blue and hyperpolarized to red and green. It differs from previously reported turtle ganglion cells in being color opponent in the entire field, not only in the surround, and in showing spatial opponency.

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