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K I Naka

Publications and source records attributed to K I Naka.

At least 19 recordsLinked to original sources

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

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

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

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

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

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

Spatial distribution of potential in a flat cell. Application to the catfish horizontal cell layers.

An analytical solution is obtained for the three-dimensional spatial distribution of potential inside a flat cell, such as the layer of horizontal cells, as a function of its geometry and resistivity characteristics. It was found that, within a very large range of parameter values, the potential is given by [Formula: see text] where r = rho/rho(0), z = z/rho(0), rho = (R(i)/R(m)).rho(0), delta = h/rho(0); K is a constant; J is the assumed synaptic current; rho, z are cylindrical coordinates; rho(0) is the radius of the synaptic area of excitation; h is the cell thickness; and R(i), R(m) are the intracellular and membrane resistivities, respectively. Formula A closely fits data for the spatial decay of potential which were obtained from the catfish internal and external horizontal cells. It predicts a decay which is exponential down to about 40% of the maximum potential but is much slower than exponential below that level, a characteristic also exhibited by the data. Such a feature in the decay mode allows signal integration over the large retinal areas which have been observed experimentally both at the horizontal and ganglion cell stages. The behavior of the potential distribution as a function of the flat cell parameters is investigated, and it is found that for the range of the horizontal cell thicknesses (10-50 mu) the decay rate depends solely on the ratio R(m)/R(i). Data obtained from both types of horizontal cells by varying the diameter of the stimulating spot and for three widely different intensity levels were closely fitted by equation A. In the case of the external horizontal cell, the fit for different intensities was obtained by varying the ratio R(m)/R(i); in the case of the internal horizontal cell it was found necessary, in order to fit the data for different intensities, to vary the assumed synaptic current J.

Animals

Dogfish ganglion cell discharge resulting from extrinsic polarization of the horizontal cells.

1. Ganglion cell discharges were evoked by extrinsic polarization of the horizontal cells in the retina of the smooth dogfish (Mustelus canis). Depolarization of the horizontal cell gave rise to a discharge similar to that evoked by a spot of light (centre type response) and hyperpolarization of the horizontal cell, a discharge similar to that by an annulus (surround type response).2. Procion dye injection established that the current-passing electrode was sometimes located in the external horizontal cell. Other possibilities, such as middle and internal horizontal cells, were neither confirmed nor excluded.3. Activation of ganglion cells by current was possible under completely dark-adapted conditions and for several log units above this level.4. Depolarizing current enhanced the ganglion cell response evoked by a light spot in the centre of its receptive field; hyperpolarizing current antagonized the response to the same flash.5. The results are consistent with the supposition that a potential change in the horizontal cell, irrespective of its polarity, or whether produced by light or current, spreads within a laminar layer (the S-space). The effect of the potential change is to modulate the response of bipolar cells and their input into the ganglion cell.

Animals