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K I Naka

Publications and source records attributed to K I Naka.

At least 19 recordsLinked to original sources
PubMed1997

Processing of color- and noncolor-coded signals in the gourami retina. I. Horizontal cells.

There are two types of horizontal cells, the luminosity and the chromaticity cells, in the retina of the kissing gourami, Helostoma rudolfi. Luminosity cells occupy the outermost layer proximal to the receptor terminals, whereas chromaticity cells form a layer proximal to the layer of luminosity cells. Neither type of cell has axons. Responses were evoked by light from red and green light-emitting diodes. The two stimuli were modulated either by a pulsatile or a white-noise signal. The luminosity cell always produced a hyperpolarizing response. The chromaticity cell produced a hyperpolarizing response when stimulated by only one color. However, in the presence of a steady or modulated green input, a red stimulus produced a depolarizing response. Such chromaticity cells were similar to the (spectral) biphasic chromaticity horizontal cells observed in other retinae. The depolarizing phase of the red response was produced by the balance of intensity of the two inputs, red and green. We used white-noise methodology to identify the dynamics of the horizontal cell's modulation response by taking advantage of the fact that a Wiener kernel is a measure of a cell's incremental sensitivity, which includes its response dynamics. Under all conditions, a steady state modulation response by both luminosity and chromaticity cells always was related linearly to the input modulation. The average mean square error (MSE) of the model predicted by the first-order kernel was approximately 8% for both luminosity (n = 116) and chromaticity (n = 23) cells. In some cases, the MSE was a few percent even when the peak-to-peak response amplitude was nearly 30 mV. The ratio of inputs from red and green cones to both types of horizontal cells was variable; the major input for luminosity cells came from red cones, whereas the major input for chromaticity cells came from green cones. First-order kernels generated by the major input were robust in terms of waveform in the sense that the waveform remained unchanged whether or not there was a steady or modulated illumination by the opposing color. The results reported here do not address the question of the neural circuitry that generates horizontal cell responses, in particular, the depolarizing response. However, whatever that circuitry might be, the high degree of linearity of the modulation response by both types of cell under various stimulus conditions imposes restrictions on the performance of any proposed model as well as on mechanisms that underlie the generation of the horizontal cell response.

Animals↗
PubMed1997

Processing of color- and noncolor-coded signals in the gourami retina. II. Amacrine cells.

The same set of stimuli and analytic methods that was used to study the dynamics of horizontal cells () 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.

Animals↗
PubMed1997

Processing of color- and noncolor-coded signals in the gourami retina. III. Ganglion cells.

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 (,). 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.

Animals↗
PubMed1997

Complexity and frequency hierarchies in the catfish retina.

The intricate connectivity and interactions between neurons in the vertebrate retina have made their individual roles in signal processing very difficult to elucidate. We have used a recently developed mathematical tool, fast orthogonal search (FOS), to probe the catfish outer (distal) and inner (proximal) retina, and study the signal processing within. Through FOS, a given waveform can be decomposed into a parsimonious sinusoidal series containing the most significant constituent frequencies. In particular, we examined the light-evoked first-order Wiener kernels of horizontal cells and on-bipolar cells, and on-off, off- and on-amacrine and ganglion cells. Here we report a hierarchy (correlation coefficient up to 0.86) in preferred frequency and complexity of response corresponding to the retina's structural hierarchy. In addition, clear differences between on-, on-off and off-cell functional characteristics were detected. For example, the kernel waveform for the on-amacrine cell was found to be more complex and to have a higher preferred frequency than that for the off-amacrine cell. Indeed FOS analysis revealed that both off- (sustained) amacrine and off-ganglion cells exhibit significantly less complexity in their waveforms for signal processing of light input than do the corresponding on- and on-off cells. This shows a clear breakdown in symmetry between on- and off-pathways, and suggests that connections to off-cells may provide fewer or a smaller variety of inputs than those to on- and on-off cells. Many of our new findings can be appreciated by assuming an underlying cascade structure for the retinal information processing. The FOS findings in particular support the following previously advanced hypothesis: the transition in nonlinear processing from on-off amacrine to on- off-amacrine cells is due to high-pass linear filtering. Furthermore, our results indicate that the high-pass filtering is more sharply differentiating for the on-amacrine than for the off-amacrine cell.

Animals↗
PubMed1990

Dissection of the neuron network in the catfish inner retina. IV. Bidirectional interactions between amacrine and ganglion cells.

1. We have functionally dissected the neuron network in the catfish inner retina by means of current injection. Simultaneous intracellular recordings were made from two neighboring neurons with the use of two separate electrodes. Extrinsic current was injected into one neuron, and the resulting intracellular responses were recorded from the other neuron. The test signals included 1) a single-frequency sinusoid, 2) a depolarizing or a hyperpolarizing current pulse, and 3) white-noise modulated current from which Wiener kernels were computed by an input-output cross-correlation process. 2. Extrinsic current injected into an ON amacrine (NA) cell evoked responses from a neighboring ON ganglion (GA) cell. Conversely, current injected into a GA cell elicited responses from a neighboring NA cell. Similar results were obtained for the transmission between OFF amacrine (NB) and OFF ganglion (GB) cells. Neural filters for the forward and backward transmissions between amacrine and ganglion cells of the same response polarity were low-pass, constant gain with a cutoff frequency of 40-50 Hz. The gain measured by current-amplitude relationships was comparable for the forward (N----G) and backward (G----N) transmission. 3. Similar bidirectional signal transmission was found between amacrine cells and between ganglion cells of the same response polarity. Neural filters for such transmission were also low-pass, constant gain with a cutoff frequency of 40-50 Hz. 4. Because a large portion of the current-evoked response was predicted by the first-order kernel, transmission between cells of the same response polarity was approximately linear. The current-evoked first-order kernels were brief and impulse-like compared with the light-evoked first-order kernels. 5. We conclude that ON and OFF amacrine and ganglion cells form two ON- and OFF-cell clusters in which cells are extensively and bidirectionally interconnected, enhancing the response in each cluster.

Animals↗
PubMed1990

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

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.

Animals↗
PubMed1987

Dendritic morphology of indoleamine cells revealed by intracellular injection of lucifer yellow in fixed carp retina.

The dendritic morphology of indoleamine amacrine cells in carp retina was investigated by identifying their fluorescent cell bodies by preloading with noradrenaline followed by iontophoretic injection of Lucifer Yellow in isolated and aldehyde-fixed preparations under microscopic control. Although two subpopulations of serotonin-like immunoreactive amacrine cells (small and large in soma size) were found, small cells were not seen in aldehyde-fixed preparations. Cells preloaded with noradrenaline corresponded to large immunoreactive cells and were labeled with Lucifer Yellow. The cell bodies labeled were located at the innermost level of the inner nuclear layer, and gave rise to three to five primary dendrites which branched frequently and were found mainly in sublamina a of the inner plexiform layer. These cells examined in an intermediate region between the optic disc and the retinal periphery were pyriform in soma shape while dendritic fields were found or oval covering an area of 0.18 +/- 0.05 mm2 (510 +/- 80 microns in diameter). Cell density in this region was about 32 cells/mm2 and, therefore, their dendritic field coverage was approximately 6.0.

Animals↗
PubMed1987

Dynamics of turtle cones.

The response dynamics of turtle photoreceptors (cones) were studied by the cross-correlation method using a white-noise-modulated light stimulus. Incremental responses were characterized by the kernels. White-noise-evoked responses with a peak-to-peak excursion of greater than 5 mV were linear, with mean square errors of approximately 8%, a degree of linearity comparable to the horizontal cell responses. Both a spot (0.17 mm diam) and a large field of light produced almost identical kernels. The amplitudes of receptor kernels obtained at various mean irradiances fitted approximately the Weber-Fechner relationship and the mean levels controlled both the amplitude and the response dynamics; kernels were slow and monophasic at low mean irradiance and were fast and biphasic at high mean irradiance. This is a parametric change and is a piecewise linearization. Horizontal cell kernels evoked by the small spot of light were monophasic and slower than the receptor kernels produced by the same stimulus. Larger spots of light or a steady annular illumination transformed the slow horizontal cell kernel into a fast kernel similar to those of the receptors. The slowing down of the kernel waveform was modeled by a simple low-pass circuit and the presumed feedback from horizontal cells onto cones did not appear to play a major role.

Animals↗
PubMed1984

Effects of dopamine on photopic L-type S-potentials in the catfish retina.

Photopic L-type responses were recorded from the soma and the axon terminal of horizontal cells of the catfish (Ictalurus punctatus) retina in eye-cup preparations. The responses were produced by a spot of light with 100-micron diameter (intensity, 10 microW/cm2), which was flashed or steadily illuminated and swept along a 6-mm length over the retinal surface at a speed of 0.95 mm/sec. In some experiments, a traveling random bar stimulus was used instead of the sweeping spot. While recording the responses, dopamine (DA) was applied in a jet form via a nebulizer over the retinal preparation or as superfusate to the eye-cup preparation. DA increased the response amplitude by about 50% and markedly narrowed the spatial profile of the responses from the soma but not from the axon terminal. These DA effects were observed in both normal retinas and those from which DA cells had been deprived by prior intraocular injection of 6-hydroxydopamine. Deprival of DA cells from the retina resulted in a slightly wider spatial profile of the soma and axon terminal responses than that in normal retinas. The results indicate that the spatial properties of photopic L-type responses are modulated by DA at the soma level of horizontal cells.

Animals↗
PubMed1983

Regional difference in density of monoamine-accumulating cells of carp and catfish retinas.

By means of a histofluorescence technique, a comparative study was conducted on the regional density of dopaminergic (DA) and indoleamine-accumulating (IA) cells in carp (Cyprinus carpio) and catfish (Ictalurus punctatus) retinas. In order to enhance detection of fluorescent cells, noradrenaline (NA; 5.0 micrograms) or a mixture of NA (2.5 micrograms) and 5,6-dihydroxytryptamine (5,6-DHT; 2.5 micrograms) was intravitreally injected into the eyes 2-3 hr before enucleation. DA and IA cells were counted systematically in space on flat-mounted preparations. Both classes of cells were found to be distributed similarly in the two species of fish; the cell density is highest in the circumferential margin of the retina, and is slightly higher in a region dorsal to the optic disc than in the surrounding area. Differences in the distribution pattern of the cells between carp and catfish retinas were as follows: (a) the DA cell density is higher over the whole retinal field in carp (the mean density +/- SD = 34 +/- 16 cells/mm2) than in catfish (13 +/- 7 cells/mm2); (b) the region where the density is slightly higher than in the surrounding area is restricted to a small area immediately dorsolateral to the optic disc in carp, while it is relatively broadly placed dorsal to the optic discs, forming a horizontal band in catfish; (c) the density ratio of DA cells to IA cells is 1:1 in carp but 1:2 in catfish; and (d) catfish DA cells seem to be more irregular than carp DA cells in shape, size, dendritic arborization, uptake preference for monoamines intravitreally injected, and also in depth location seen in radial cryosections.

Animals↗
PubMed1983

Synaptic organization involving receptor, horizontal and on- and off-center bipolar cells in the catfish retina.

HRP-injected bipolar cells were analyzed electron microscopically in catfish retina. The dendrites of the two types of bipolar cells (on-center and off-center) invaginated into the photoreceptor terminals in distinct patterns. Those from off-center bipolars ended as the central element of triads in such a way that every synaptic ridge examined was contacted by a single process along its apex. In contrast, labeled processes from on-center bipolars never made direct contact with the synaptic ridges of any photoreceptors. In addition, chemical synapses made by external horizontal cells onto photoreceptors were observed. The postsynaptic sites was always a telodendrial process of a photoreceptor.

Animals↗
PubMed1981

Direct electrical connections between transient amacrine cells in the catfish retina.

Transient amacrine cells were identified by their intracellularly recorded response to flashes of light. These cells typically respond with a transient depolarization, often followed by a steady-state response during the stimulus. When two electrodes were placed in different transient amacrine cells, current of either polarity passed through one electrode produced a steady-state voltage change that was recorded by the electrode in the nearby cell. Following identification of the physiological type, transient amacrine cells were injected with horseradish peroxidase and the tissue was processed for light and electron microscopy. Both conventional chemical synaptic junctions and gap junctions were found to connect amacrine cells.

Animals↗
PubMed1979

Adaptation in catfish retina.

1. We define absolute sensitivity as (voltage/illuminance) and incremental sensitivity as the peak-to-peak amplitude of the first-order (Wiener) kernels. 2. Incremental sensitivity of the horizontal cells is the local slopes of the Michaelis-Menten equation and that of more proximal neurons is the Fechner slope. In a log-log plot, the former has a slope of -2, whereas the latter a slope of -1, as predicted by Williams and Gale (39). 3. During a moderate to strong steady illumination, absolute sensitivity decreases but incremental sensitivity increases. The reverse occurs during dark adaptation. 4. The presence of a steady illumination did not prevent signal transmission from horizontal to ganglion cells. 5. From these results we conclude that: adaptation in the catfish retina includes two components: a) a lateral shift of the voltage-intensity curve along the intensity axis, and b) changes in the time course of light-evoked response. We argue that the latter phenomenon is related to the presumed horizontal cell-to-receptor cell negative feedback.

Adaptation, Physiological↗
PubMed1978

gamma-Aminobutyric acid: a neurotransmitter candidate for cone horizontal cells of the catfish retina.

In the catfish retina, horizontal cells that receive inputs exclusively from red-sensitive cones are the only neurons that accumulate exogenous gamma-aminobutyric acid under our experimental conditions. When isolated eyecups are perfused with bicuculline methochloride, an antagonist of postsynaptic gamma-aminobutyric acid receptors, responses of cone photoreceptors to a field of light (3 mm in diameter) become much slower and approach those to a small spot of light (0.3 mm). In addition, bicuculline methochloride decreases the frequency responses of cone horizontal cells to a field of light. These findings indicate that, in the catfish retina, feedback synapses from cone horizontal cells to cones are chemically mediated and may use gamma-aminobutyric acid as a neurotransmitter. Our results also confirm the hypothesis that, in the catfish retina, a function of the negative feedback is to improve the frequency responses of the system.

Action Potentials↗