Horizontal cell sensitivity in the cat retina during prolonged dark adaptation
Lankheet, M.J.M. ; Rowe, M.H. ; Wezel, R.J.A. Van; De Grind, W.A. Van - \ 1996
Visual Neuroscience 13 (1996)5. - ISSN 0952-5238 - p. 885 - 896.
Cat retina - Dark adaptation - Horizontal cell - Rod and cone input
The effects of dark adaptation on the response properties of ganglion cells have been documented extensively in the cat retina. To pinpoint the different retinal mechanisms that underlie these effects, we studied the response characteristics of cat horizontal (H) cells during prolonged dark adaptation. H-cell responses were recorded intracellularly in the optically intact, in vivo eye. To disentangle rod and cone contributions, sensitivity changes during dark adaptation were tracked with white light and with monochromatic lights that favored either rod or cone excitation. Stable, long-lasting recordings allowed us to measure changes of sensitivity for adaptation periods up to 45 min. Thresholds for white light and 503-nm monochromatic light decreased steadily and in parallel. The maximum increase of sensitivity, after extinguishing a photopic adaptation light, was 1.8 log units only, reached after about 35 min. Sensitivity for 581-nm lights also increased steadily, but at a shallower slope. The steady increase of sensitivity was concomitant with a linear shift in resting membrane potential and with an increase in relative rod contribution to the threshold responses. Even though small-amplitude responses were rod dominated after prolonged dark adaptation, sensitivity to rod signals remained relatively low, compared to sensitivity of cone responses or to the absolute sensitivity of ganglion cells. This suggests that the cone-H-cell pathway plays no role in the dark-adapted cat retina.
Gain control and hyperpolarization level in cat horizontal cells as a function of light and dark adaptation
De Grind, W.A. Van; Lankheet, M.J.M. ; Wezel, R.J.A. Van; Rowe, M.H. ; Hulleman, J. - \ 1996
Vision Research 36 (1996)24. - ISSN 0042-6989 - p. 3969 - 3985.
Cat retina - Dark adaptation - Horizontal cells - Light adaptation - Rod-cone interaction
First a model is presented that accurately summarizes the dynamic properties of cat horizontal (H-) cells under photopic conditions as measured in our previous work. The model predicts that asymmetries in response to dark as compared to light flashes are flash-duration dependent. This somewhat surprising prediction is tested and confirmed in intracellular recordings from the optically intact in vivo eye of the cat (Experiment 1). The model implies that the gain of H-cells should be related rather directly to the sustained (baseline) membrane potential. We performed three additional experiments to test this idea. Experiment 2 concerns response vs intensity (R-I-) curves for various flash-diameters and background-sizes with background luminance varying over a 4 log unit range. Results support the assumption of a rather strict coupling between flash sensitivity (gain) and the sustained level of hyperpolarization. In Experiment 3 we investigate this relation for both dark and light flashes given on each of four background light levels. The results suggest that there are fixed minimum and maximum hyperpolarization levels, and that the baseline hyperpolarization for a given illumination thus also sets the available range for dark and light flash-responses. The question then arises whether, or how this changes during dark adaptation, when the rod contribution to H-cell responses gradually increases. The fourth experiment therefore studies the relationship between gain and hyperpolarization level during prolonged dark-adaptation. The results show that the rod contribution increases the polarization range of H-cells, but that the gain and polarization level nevertheless remain directly coupled. H-cell models relying on a close coupling between polarization level and gain thus remain attractive options.
Spatial and temporal properties of cat horizontal cells after prolonged dark adaptation
Lankheet, M.J.M. ; Rowe, M.H. ; Wezel, R.J.A. Van; De Grind, W.A. Van - \ 1996
Vision Research 36 (1996)24. - ISSN 0042-6989 - p. 3955 - 3967.
Cat retina - Dark adaptation - Horizontal cell - Spatial and temporal properties
We studied the change of spatial and temporal response properties for cat horizontal (H-) cells during prolonged dark adaptation. H-cell responses were recorded intracellularly in the optically intact, in vivo eye. Spatial and temporal properties were first measured for light-adapted H-cells, followed by a period of dark adaptation, after which the same measurements were repeated. During dark adaptation threshold sensitivity was measured at regular intervals. Stable, long lasting recordings allowed us to measure changes of sensitivity and receptive field characteristics for adaptation periods up to 45 min. Although cat H-cells showed no signs of dark suppression or light sensitization, they remained insensitive in the scotopic range, even after prolonged dark adaptation. Absolute thresholds were in the low mesopic range. The sensitization was brought about by a shift from cone to rod input, and by substantial increases of both spatial and temporal integration upon dark adaptation. The length constant in the light-adapted state was on average about 4 deg. After dark adaptation it was up to a factor of three larger, with a median ratio of 1.85. Response delays, latencies and durations for (equal amplitude) threshold flash responses substantially increased during dark adaptation.
Nonlinearity and oscillations in X-type ganglion cells of the cat retina
Przybyszewski, A.W. ; Lankheet, M.J.M. ; De Grind, W.A. Van - \ 1993
Vision Research 33 (1993)7. - ISSN 0042-6989 - p. 861 - 875.
Cat retina - Intracellular ganglion cell recordings - Transient oscillations - van der Pol oscillator
Intracellularly recorded light-responses of X-type ganglion cells in the cat retina were separated, with the help of a wavelet method, into "slow" membrane ("G")-potentials and the corresponding spike trains. In response to sinusoidally modulated high intensity light spots with different sizes and frequencies, X-type ganglion cells show both oscillations correlated with the stimulus frequency and other, faster, oscillations that were not always locked to the stimulus. A forced van der Pol oscillator model with stimulus-dependent coefficients proved to describe the empirical findings quite well. A linearity-coefficient of the equations indicates strong nonlinearity at a temporal frequency of 8 Hz with spot sizes on the order of 0.5-0.7 deg and decreasing nonlinearity at lower temporal frequencies or smaller spot sizes, while the faster oscillations become more prominent. We could not determine whether the oscillations are intrinsic to the cell-membrane or generated by (or in interaction with) the preganglionic retinal meshwork. The results show that X-cell spike-trains can contain oscillations that are not phase-locked to the stimulus and that are therefore virtually invisible after stimulus synchronous averaging. It is not likely that these retinal oscillations directly induce the well described oscillations in cat visual cortex, since they usually fall in a different frequency range.
Responses of cat horizontal cells to sinusoidal gratings
Lankheet, M.J.M. ; Prickaerts, J.H.H.J. ; Grind, W.A. van de - \ 1992
Vision Research 32 (1992)6. - ISSN 0042-6989 - p. 997 - 1008.
Cat retina - Horizontal cell - Sine grating - Spatiotemporal transfer
The spatiotemporal properties of cat horizontal (H-) cells were studied by recording the intracellular responses in the optically intact, in vivo, eye to sinusoidal gratings at a photopic mean illumination level. In order to investigate the linearity of spatial summation a "null test" was performed in which the responses to contrast reversal gratings were measured at different positions of the grating relative to the receptive field. Spatial and temporal transfer functions were measured using drifting sinusoidal gratings of variable spatial and temporal frequencies. The amplitudes of cat H-cell responses to contrast reversal gratings modulated with a square wave time-course showed a sinusoidal dependence on spatial phase. When zero crossings of the grating were lined up with the receptive field center, as defined by the maximum of the measured line weighting function, contrast reversal produced no response modulation. This result did not depend on the spatial frequency of the grating or the temporal frequency of contrast modulation over substantial ranges. The response waveform was found not to depend on the spatial phase of the grating. The spatial transfer function of cat H-cells has low-pass characteristics with a cut-off frequency in the range of about 0.4-1.5 c/deg. The shape of the spatial transfer function was roughly the same for temporal frequencies ranging from 3 to 10 Hz. The temporal transfer function exhibited band-pass characteristics with a maximum response amplitude at 3-6 Hz. The amplitude fall-off for low and high temporal frequencies was independent of the spatial frequency of the grating. The results obtained with sine gratings were found not to agree with the receptive field profiles measured with narrow slits flashed at different positions in the receptive field.
Effects of background illumination on cat horizontal cell responses
Lankheet, M.J.M. ; Wezel, R.J.A. van; Grind, W.A. van de - \ 1991
Vision Research 31 (1991)6. - ISSN 0042-6989 - p. 919 - 932.
Cat retina - Horizontal cells - Light adaptation
The process of light adaptation in cat horizontal cells was studied by means of intracellular recordings in the optically intact, in situ eye. Response vs intensity profiles were measured using increment- as well as decrement-flashes of "white light" on steady backgrounds. The effects of background illumination in the mesopic and photopic range on both purely rod-driven horizontal cells and mixed rod and cone input horizontal cells were investigated. Increasing the background illumination for mixed-input horizontal cells strongly reduced the contribution from the rod system. The rod aftereffect in the responses to high intensity flashes is totally suppressed at higher background levels. Light adaptation resulted in a sustained hyperpolarization without a substantial effect on the total response range of the response vs intensity curve. At higher background intensities more of the response range is made up of depolarizing responses to decrements of light. Increasing the background illumination also shifted the operating curve to higher intensities. Increment threshold functions, measured with a 3.9 deg diameter test spot on a large background (8.8 deg diameter) showed a linear relation between log threshold intensity and log background intensity with a slope of, on average, 0.64. The response vs intensity curve for the rod horizontal cell typically spanned a narrower intensity range and was displaced toward lower intensities as compared to that for the mixed input horizontal cells. Background illumination greatly reduced the total response range for rod horizontal cells. Increment threshold curves for rod horizontal cells clearly indicated loss in sensitivity due to response compression.