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Comparative aspects of horizontal ocular reflexes and their cerebellar adaptive control in vertebrates.

1. Vestibuloocular reflex and optokinetic eye movement response are ocular reflexes conjointly subserving stability of retinal images. 2. These reflexes exhibit marked adaptability under sustained imposition of retinal errors. 3. Comparative studies in various vertebrate species reveal the importance of this adaptability as fundamental neural function based on plasticity of cerebellar synapses.

Animals↗

Hard contact lenses alter accommodative gain but do not prevent refractive adaptation in chicks.

This study compared the compensatory response to hyperopic defocus imposed on chicks in two different ways: (1) with-10 D spectacle lenses, and (2) with plano hard contact lens. The hyperopia seen with the contact lenses in situ was a consequence of their flat profile relative to the chick cornea, resulting in a negative fluid lens of approximately 16 D at day 2 and 9 D by day 10. This decrease with age reflects the corneal flattening that accompanies normal eye growth. By optically neutralizing the cornea, the contact lenses also had two other important effects: (1) a reduction in refractive astigmatism to almost negligible levels, and (2) a reduction in accommodative gain. The latter effect reflects the loss of the corneal component of the chick's accommodation and was estimated to be of the order of 40 to 57%, based on measurements made using topically applied nicotine to stimulate accommodation. Thus any estimate of the imposed hyperopic defocus based on accommodative effort required to overcome such errors will be too large. Chicks wearing either lens type on a continuous basis from hatching to 10 days only partially compensated for the imposed hyperopia through an increase in vitreous chamber growth. However, the effects were smaller in the spectacle lens group (e.g., a mean myopic shift of -4.1 +/- 2.3 D compared to -6.3 +/- 2.4 D for the contact lens group at day 10), although both groups experienced similar amounts of hyperopic defocus around day 10 (effective power of -10 D spectacle lens: -9.4 D). The changes seen in the spectacle lens group thus represent poorer compensation, i.e., 44 vs. 71% of the imposed error. However, overcompensation is the predicted effect, if any, of the accommodative deficit imposed on the contact lens group, and this was not seen. That compensation, albeit incomplete, occurred with the contact lens as well as the spectacle lens, suggests that neither accommodation nor astigmatism are fundamental cues for emmetropization as modeled here.

Accommodation, Ocular↗

Changes in contrast sensitivity induced by defocus and their possible relations to emmetropization in the chicken.

PURPOSE: To test whether the level of contrast adaptation (CA) relates to refractive development in the chicken. (CA refers to a spatial frequency-selective increase of suprathreshold contrast sensitivity after exposure to low-contrast patterns). METHODS: CA was determined in individual chicks by comparing their optomotor gain in response to drifting low-contrast stripe patterns before and after treatment with spectacle lenses. The amount of CA was compared with the loss of contrast predicted from defocus at the tested spatial frequency. The reversion of CA and recovery from deprivation myopia were studied while the retinal image features were controlled by forcing the animals to watch spatially filtered digital video clips. RESULTS: CA was induced by wearing positive and negative lenses for 1.5 hours, both without and with cycloplegia, but was less pronounced in the case of positive lenses when accommodation was intact. The amount of CA at a tested spatial frequency was predicted from the loss of contrast calculated from the modulation transfer function for a defocused optical system. Watching low-pass-filtered video clips induced deprivation myopia and inhibited recovery from it. It also prevented the reversal of CA that was previously induced by deprivation. Both recovery from deprivation myopia and recovery from CA occurred with sharp video clips, although less so than with normal visual exposure. CONCLUSIONS: CA changes with retinal image sharpness and occurs even when accommodation is intact. Because CA correlates with myopia induced by frosted occluders, negative lenses, and low-pass-filtered video clips, and its reversal correlates with recovery from myopia, it is possible that shifts in CA may represent a signal related to refractive error development.

Accommodation, Ocular↗

Dark- and light-induced changes in coupling between horizontal cells in mammalian retina.

Retinal horizontal cells exhibit large receptive fields derived from their extensive electrical coupling by means of gap junctions. The conductance of these gap junctions seems to be regulated by dopamine acting through a cAMP-mediated cascade. There is now abundant evidence that extracellular dopamine levels vary with changes in ambient light intensity, suggesting that changes in the dark/light adaptational state of the retina can modulate coupling between horizontal cells. We studied this question in the mammalian retina by determining the effects of ambient light levels, in the form of changing background light intensity, on the coupling profiles of A- and B-type horizontal cells in the rabbit. Changes in coupling were assessed by measurements of the space constants of the syncytium formed by horizontal cells and the intercellular spread of the biotinylated tracer Neurobiotin. Our results indicate that dark-adapted horizontal cells show relatively weak coupling. However, presentation of background lights as dim as one-quarter log unit above rod threshold resulted in increases in both the averaged extent of tracer coupling and space constants of A- and B-type horizontal cells. Coupling expanded further as background light intensities were increased by 1-1.5 log units, after which additional light adaptation brought about an uncoupling of cells. Coupling reached its minimum at light intensities about 3 log units above rod threshold, after which, with further light adaptation, it stabilized at levels close to those seen in dark-adapted retinas. Our results indicate that electrical coupling between mammalian horizontal cells is modulated dramatically by changes in the adaptational state of the retina: coupling is maximized under dim ambient light conditions and diminishes as the retina is dark or light adapted from this level.

Adaptation, Ocular↗

Rod and cone contribution to adaptation processes in cat retinal ganglion cells.

Extracellular ganglion cell responses were recorded to investigate mechanisms of light adaptation. Monochromatic test spots (575 nm) were projected onto the receptive field center of off-center cells and superimposed on a steady blue-green Ganzfeld background (Schott Filter BG 28), the strength of which was increased in steps of 0.5 log units to adapt rods. Response vs. log intensity functions were determined over a range of 7 log units of test light irradiance at each background level. At higher adaptation levels response thresholds followed the typical Weber function. Surprisingly at lower adaptation levels the sensitivity of the cell increased by about 0.7 log units, most markedly in a range of 1 log unit of moderate light adaptation when the background was changed from dark to the dimmest detectable background (10(-5) lm/m2). In the dark-adapted state a small off-response of long latency (40-100 ms at 10(2) quanta.s-1.microns-2) is observed at low rod stimulating test light irradiances. A transition to a cone-dominated transient response of 2 to 5 ms duration occurred at high intensities (10(5) quanta.s-1.microns-2). At mesopic levels the two responses seem to cancel each other, rendering a delayed off-response that is probably the result of rod-cone interaction. As in psychophysics, saturation can be observed at very high background intensities (10(6) quanta.s-1 microns-2). These data suggest interactions between rods and cones that determine the sensitivity of cat retinal ganglion cells at low levels of adaptation for suprathreshold stimuli.

Action Potentials↗

Adaptation and dynamics in X-cells and Y-cells of the cat retina.

On the basis of the spatial summation properties of their receptive fields, cat retinal ganglion cells were classified as either X-cells (linear) or Y-cells (non-linear). Responses were then obtained to a small, centered spot, square-wave modulated in time and superimposed on various levels of diffuse, steady background illumination. When fully dark-adapted, both X-cells and Y-cells produced responses that were entirely sustained. When well light-adapted but still in the scotopic range, both cell types produced largely transient responses with only a very small sustained component. The sustained or transient nature of responses is, therefore, not an invariant characteristic of X-cells and Y-cells in the scotopic range. We also conclude that the mechanism which controls the center's sensitivity in the scotopic range is similar though not identical in the two types of cells.

Adaptation, Ocular↗

[The alteration of visual evoked potentials in light- and dark-adapted states as a measure for the visual field sensibility at elevated intraocular pressure (author's transl)].

On 12 voluntary subjects with normal visual function the intraocular pressure was increased dependent on the ophthalmic blood pressure Pm = (Ps + Pd)/2. The visual field was tested under this pressure load by means of a subjective method and the visual evoked potential was derived under light and dark adaptation. Comparison of subjective and objective investigations demonstrated that the different visual evoked potentials which are dependent on the state of adaptation are a measure for the pressure sensibility of prelaminar optic nerve fibers.

Adaptation, Ocular↗

An analysis of rod outer segment adaptation based on a simple equivalent circuit.

This model of rod outer segment adaptation is based on the hypothesis that transmitter substance released by bleached rhodopsin closes sodium channels in the outer segment plasma membrane, leading to hyperpolarization of the receptor. The outer segment adaptation processes of the model are associated with the transmitter release, the transmitter background concentration and the plasma membrane leakage. Changes in the three model parameters correspond to the three types of outer segment adaptation process. According to the model the stimulus-response function is in every adaptational state U/Umax = I/(I + IH). The model predicts how each adaptation process affects IH and Umax. Specifically, when the number of liberated transmitter molecules per isomerizing quantum decreases, the amplitude Umax remains constant but IH increases. A short description of this model has been published in a paper reporting experimental results on background adaptation (Hemilä, 1977).

Adaptation, Ocular↗