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M Sakuranaga

Publications and source records attributed to M Sakuranaga.

10 recordsLinked to original sources

Dynamics of skate horizontal cells.

The all-rod retina of the skate (Raja erinacea or R. oscellata) is known to have the remarkable capability of responding to incremental flashes superimposed on background intensities that initially block all light-evoked responses and are well above the level at which rods saturate in mixed rod/cone retinas. To examine further the unusual properties of the skate visual system, we have analyzed responses of their horizontal cells to intensity-modulated step, sinusoidal, and white-noise stimuli. We found that during exposures to mean intensities bright enough to block responses to incremental stimuli, decremental stimuli were also initially blocked. Thereafter, the horizontal cells underwent a slow recovery phase during which there was marked nonlinearity in their response properties. The cell first (within 2-3 min) responded to decrements in intensity and later (after greater than 10 min) became responsive to incremental stimuli. After adaptation to a steady state, however, the responses to intensity modulation were nearly linear over a broad range of modulation depths even at the brightest mean levels of illumination. Indeed, examination of the steady-state responses over a 5-log-unit range of mean intensities revealed that the amplitude of the white noise-evoked responses depended solely on contrast, and was independent of the retinal irradiance as the latter was increased from 0.02 to 20 muW/cm2; i.e., contrast sensitivity remained unchanged over this 1,000-fold increase in mean irradiance. A decrement from the mean as brief as 2 s, however, disturbed the steady state. Another unexpected finding in this all-rod retina concerns surround-enhancement, a phenomenon observed previously for cone-mediated responses of horizontal cells in the retinas of turtle and catfish. While exposure to annular illumination induced response compression and a pronounced sensitivity loss in response to incremental light flashes delivered to the dark central region, the cell's sensitivity showed a significant increase when tested with a white noise or sinusoidally modulated central spot. Unlike horizontal cells in other retinas studied thus far, however, response dynamics remained unchanged. Responses evoked either by a small spot (0.25-mm diam) or by a large field light covering the entire retina were almost identical in time course. This is in contrast with past findings from cone-driven horizontal cells whose response waveform (dynamics) was dependent upon the size of the retinal area stimulated.

Animals

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.

Animals

Nonlinear analysis: mathematical theory and biological applications.

Wiener's method of a nonlinear system analysis and its application to neurophysiology are surveyed. His theory is explained on the orthogonal functional series expansion of a nonlinear noise with respect to the Brownian motion whose formal derivative is understood as the white Gaussian noise. Then, efforts made for applying the method to practice are discussed. Special attention is paid on relationship between the internal structure of a given nonlinear system and its kernels of the functionals. This serves to explain a meaning of the kernels. Finally, following the works of one of the authors (K-I.N.) and his colleagues, the analysis of the catfish retinal neuron system is introduced as a typical and successful example of the applications in neurophysiology.

Adaptation, Ocular

Visual sensitivity and Wiener kernels.

Visual sensitivity is defined in terms of the static and dynamic parts of a photo-evoked response. The first-order kernel induced by a white-noise modulated light is associated directly with both the incremental and contrast sensitivities classically defined, making a comprehensive measure of neuron sensitivity in the visual system.

Humans

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.

Animals

Signal transmission in the catfish retina. I. Transmission in the outer retina.

Extrinsic current, either pulsatile or white-noise modulated, was injected into the (cone) horizontal-cell soma and axon, and resulting responses were recorded from nearby points. In the case of white-noise inputs, signal transmission between the two points was characterized by Wiener kernels. The signal transmission within the lamina, the S-space, formed by the (cone) horizontal-cell somas and axons is quasi-linear and very fast, indicating that the laminae are purely resistive networks within the frequency range of the light-evoked response. There exists signal transaction between the lamina formed by the somas and axons. The forward transmission is constant gain, low pass, but there is a filter for the reverse transmission to impede the backflow of high-frequency components. Signals in the horizontal-cell soma are transmitted to the bipolar cells. The transmission is sign noninverting for the on-center bipolar cells and sign inverting for the offcenter cells. The transmission is quasi-linear excluding complex mechanisms in the transmission. We believe that the forward and direct transmission of signals from the horizontal to bipolar cells is the most straightforward interpretation of the observation. The transfer functions between the horizontal and bipolar cells differ considerably from one bipolar cell to the next.

Animals

Signal transmission in the catfish retina. II. Transmission to type-N cell.

Responses from channel catfish type-N (sustained amacrine) cells were evoked either by step changes in illuminance, i.e. brightening or dimming from a mean illuminance, or by a white-noise modulated light stimulus. Current injected into the horizontal-cell soma or axon produced responses in type-N cells that were very similar to those produced by light stimuli. Light- and current-evoked responses had linear and second- and third-order nonlinear components; the former contributed 40-50%, whereas the latter contributed 20-30% to the total response. The remainder of the response could have been due to higher-order nonlinearities or to intrinsic as well as extrinsic noise. Nonlinear components in the light- and current-evoked responses were sharp transient peaks, which were prominent in white-noise-evoked responses, and oscillatory wavelets. The high-frequency components in the cell's response, which result from nonlinearity, were absent in the responses from bipolar and horizontal cells. The nonlinear responses were predicted by the second- and third-order kernels. The type-N cell response was complex because the response had both linear and nonlinear components, and because of the complexities of second- and, probably, third-order kernels. The cell's complex response reflects the complex nature of the cell's function as well as its synaptic organization.

Animals

Signal transmission in the catfish retina. III. Transmission to type-C cell.

Current injected into horizontal-cell somas and axons produced transient (on-off) depolarizations from type-C cells (commonly known as transient amacrine cells) similar to those produced by light. Both the light- and current-induced responses had very small linear components and nonlinear components as represented by the second-order kernels, which reproduced the cell's response with a reasonable degree of accuracy. The second-order kernels were well defined and stereotyped. The quadratic nature of the nonlinear component is reflected in the frequency doubling response as well as the very steep input-output relationship of the cell. Type-C cell's responses evoked by light and current differed in a subtle but distinct fashion, and this difference appeared in the signature of the second-order kernels. The light-produced kernels had two diagonal positive peaks and off-diagonal valleys ("four-eye" structure), whereas the current-produced kernels had a single on-diagonal positive peak with off-diagonal negative valleys ("three-eye" structure). The difference in the kernel configuration was reflected in the cell's step-evoked response. Some type-C cells produced faster responses whereas others produced slower responses, whether evoked by light or by current. Our past and present results show that type-C cells produce a very nonlinear response that is not necessarily complex.

Animals