Frances J. Rucker

Cone signals for spectacle-lens compensation: Differential responses to short and long wavelengths

Chick eyes compensate for defocus imposed by spectacle lenses by making compensatory changes in eye length and choroidal thickness, a laboratory model of emmetropization. To investigate the roles of longitudinal chromatic aberration and of chromatic mechanisms in emmetropization, we examined the participation of different cone classes, and we compared the efficacy of lens compensation under monochromatic illumination with that under white light of the same illuminance to the chick eye. Chicks wore positive or negative 6D or 8D lenses on one eye for 3 days, under either blue (460 nm) or red (620 nm) light at 0.67 lux or under white light at 0.67 or 0.2 lux (all measures are corrected for chick photopic sensitivity). The illumination conditions were chosen to differentially stimulate either the short-wavelength and ultraviolet cones or the long-wavelength and double cones. Measurements are expressed as the relative change: the inter-ocular difference in the amount of change over the 3 days of lens wear. We find that under this low illumination the two components of lens compensation were differentially affected by the monochromatic illumination: in blue light lens compensation was mainly due to changes in eye length, whereas in red light lens compensation was mainly due to changes in choroidal thickness. In general, white light produced better lens compensation than monochromatic illumination. NEGATIVE LENSES: Under white light negative lenses caused an increase in eye length (60 microm) together with a decrease in choroidal thickness (-51 microm) relative to the fellow eye. Under blue light, although there was an increase in eye length (32 microm), there was no change in choroidal thickness (5 microm). In contrast, under red light there was a decrease in choroidal thickness (-62 microm) but no increase in eye length (8 microm). Relative ocular elongation was the same in white and monochromatic light. POSITIVE LENSES: Under white light positive lenses caused a decrease in eye length (-142 microm) together with an increase in choroidal thickness (68 microm) relative to the fellow eye. Under blue light, there was a decrease in eye length (-64 microm), but no change in choroidal thickness (2 microm). In contrast, under red light there was an increase (90 microm) in choroidal thickness but less of a decrease (-36 microm) in eye length. Lens compensation by inhibition of ocular elongation was less effective under monochromatic illumination than under white light (white v red: p=0.003; white v blue p=.014). The differential effects of red and blue light on the choroidal and ocular length compensatory responses suggest that they are driven by different proportions of the cone-types, implying that, although chromatic contrast is not essential for lens compensation and presumably for emmetropization as well, the retinal substrates exist for utilizing chromatic contrast in these compensatory responses. The generally better lens compensation in white than monochromatic illumination suggests that longitudinal chromatic aberration may be used in lens compensation.

Accommodation responses to stimuli in cone contrast space.

The aim was to identify the cone contributions and pathways for reflex accommodation. Twelve illumination conditions were used to test specified locations in cone-contrast space. Accommodation was monitored continuously in a Badal optometer while the grating stimulus (2.2 c/d sine-wave; 0.27 modulation) moved sinusoidally (0.195 Hz) towards and away from the eye from a mean position of 2.00 D (+/-1.00 D). Mean accommodation level and dynamic gain and phase at 0.195 Hz were calculated. Mean accommodation level varied significantly when the long- and middle-wavelength cone contrast ratio was altered in both the luminance and chromatic quadrants of cone-contrast space. This experiment indicates that L- and M-cones contribute to luminance and chromatic signals that produce the accommodation response, most likely through magno-cellular and parvo-cellular pathways, respectively. The L:M cone weighting to the luminance pathway that mediates accommodation is 1.63:1. The amplitude and direction of the response depends on changes in chromatic contrast and luminance contrast signals that result from longitudinal chromatic aberration and defocus of the image.

Cone contributions to signals for accommodation and the relationship to refractive error.

The accommodation response is sensitive to the chromatic properties of the stimulus, a sensitivity presumed to be related to making use of the longitudinal chromatic aberration of the eye to decode the sign of the defocus. Thus, the relative sensitivity to the long- (L) and middle-wavelength (M) cones may influence accommodation and may also be related to an individuals refractive error. Accommodation was measured continuously while subjects viewed a sine wave grating (2.2c/d) that had different cone contrast ratios. Seven conditions tested loci that form a circle with equal vector length (0.27) at 0, 22.5, 45, 67.5, 90, 120, 145 deg. An eighth condition produced an empty field stimulus (CIE (x,y) co-ordinates (0.4554, 0.3835)). Each of the gratings moved at 0.2 Hz sinusoidally between 1.00 D and 3.00 D for 40s, while the effects of longitudinal chromatic aberration were neutralized with an achromatizing lens. Both the mean level of accommodation and the gain of the accommodative response, to sinusoidal movements of the stimulus, depended on the relative L and M cone sensitivity: Individuals more sensitive to L-cone stimulation showed a higher level of accommodation (p=0.01; F=12.05; ANOVA) and dynamic gain was higher for gratings with relatively more L-cone contrast. Refractive error showed a similar correlation: More myopic individuals showed a higher mean level of accommodation (p<0.01; F=11.42; ANOVA) and showed higher gain for gratings with relatively more L-cone than M-cone contrast (p=0.01; F=10.83 ANOVA). If luminance contrast is maximized by accommodation, long wavelengths will be imaged behind the photoreceptors. Individuals in whom luminance is dominated by L-cones may maximize luminance contrast both by accommodating more, as shown here, and by increased ocular elongation, resulting in myopia, possibly explaining the correlations reported here among relative L/M-cone sensitivity, refractive error and accommodation.

Chicks use changes in luminance and chromatic contrast as indicators of the sign of defocus.

As the eye changes focus, the resulting changes in cone contrast are associated with changes in color and luminance. Color fluctuations should simulate the eye being hyperopic and make the eye grow in the myopic direction, while luminance fluctuations should simulate myopia and make the eye grow in the hyperopic direction. Chicks without lenses were exposed daily (9 a.m. to 5 p.m.) for three days on two consecutive weeks to 2 Hz sinusoidally modulated illumination (mean illuminance of 680 lux) to one of the following: in-phase modulated luminance flicker (LUM), counterphase-modulated red/green (R/G Color) or blue/yellow flicker (B/Y Color), combined color and luminance flicker (Color + LUM), reduced amplitude luminance flicker (Low LUM), or no flicker. After the three-day exposure to flicker, chicks were kept in a brooder under normal diurnal lighting for four days. Changes in the ocular components were measured with ultrasound and with a Hartinger Coincidence Refractometer (aus Jena, Jena, East Germany. After the first three-day exposure, luminance flicker produced more hyperopic refractions (LUM: 2.27 D) than did color flicker (R/G Color: 0.09 D; B/Y Color: -0.25 D). Changes in refraction were mainly due to changes in eye length, with color flicker producing much greater changes in eye length than luminance flicker (R/G Color: 102 μm; B/Y Color: 98 μm; LUM: 66 μm). Our results support the hypothesis that the eye can differentiate between hyperopic and myopic defocus on the basis of the effects of change in luminance or color contrast.

The effects of longitudinal chromatic aberration and a shift in the peak of the middle-wavelength sensitive cone fundamental on cone contrast

Longitudinal chromatic aberration is a well-known imperfection of visual optics, but the consequences in natural conditions, and for the evolution of receptor spectral sensitivities are less well understood. This paper examines how chromatic aberration affects image quality in the middle-wavelength sensitive (M-) cones, viewing broad-band spectra, over a range of spatial frequencies and focal planes. We also model the effects on M-cone contrast of moving the M-cone fundamental relative to the long- and middle-wavelength (L- and M-cone) fundamentals, while the eye is accommodated at different focal planes or at a focal plane that maximizes luminance contrast. When the focal plane shifts towards longer (650 nm) or shorter wavelengths (420 nm) the effects on M-cone contrast are large: longitudinal chromatic aberration causes total loss of M-cone contrast above 10-20 c/d. In comparison, the shift of the M-cone fundamental causes smaller effects on M-cone contrast. At 10 c/d a shift in the peak of the M-cone spectrum from 560 to 460 nm decreases M-cone contrast by 30%, while a 10 nm blue-shift causes only a minor loss of contrast. However, a noticeable loss of contrast may be seen if the eye is focused at focal planes other than that which maximizes luminance contrast. The presence of separate long- and middle-wavelength sensitive cones therefore has a small, but not insignificant cost to the retinal image via longitudinal chromatic aberration. This aberration may therefore be a factor limiting evolution of visual pigments and trichromatic color vision.

Accommodation with and without short-wavelength-sensitive cones and chromatic aberration

Accommodation was monitored while observers (23) viewed a square-wave grating (2.2 cycles/deg; 0.53 contrast) in a Badal optometer. The grating moved sinusoidally (0.2 Hz) to provide a stimulus between -1.00 D and -3.00 D during trials lasting 40.96 s. There were three illumination conditions: 1. Monochromatic 550 nm light to stimulate long-wavelength-sensitive cones (L-cones) and medium-wavelength-sensitive cones (M-cones) without chromatic aberration; 2. Monochromatic 550 nm light+420 nm light to stimulate long-, medium- and short-wavelength-sensitive cones (S-cones) with longitudinal chromatic aberration (LCA); 3. Monochromatic 550 nm light+420 nm light to stimulate L-, M- and S-cones viewed through an achromatizing lens. In the presence of LCA mean dynamic gain decreased (p=0.0003; ANOVA) and mean accommodation level was reduced (p=0.001; ANOVA). The reduction in gain and increased lag of accommodation in the presence of LCA could result from a blue-yellow chromatic signal or from a larger depth-of-focus.

The role of luminance and chromatic cues in emmetropisation.

At birth most, but not all eyes, are hyperopic. Over the course of the first few years of life the refraction gradually becomes close to zero through a process called emmetropisation. This process is not thought to require accommodation, though a lag of accommodation has been implicated in myopia development, suggesting that the accuracy of accommodation is an important factor. This review will cover research on accommodation and emmetropisation that relates to the ability of the eye to use colour and luminance cues to guide the responses.

Blue Light Protects Against Temporal Frequency Sensitive Refractive Changes

PURPOSE Time spent outdoors is protective against myopia. The outdoors allows exposure to short-wavelength (blue light) rich sunlight, while indoor illuminants can be deficient at short-wavelengths. In the current experiment, we investigate the role of blue light, and temporal sensitivity, in the emmetropization response. METHODS Five-day-old chicks were exposed to sinusoidal luminance modulation of white light (with blue; N = 82) or yellow light (without blue; N = 83) at 80% contrast, at one of six temporal frequencies: 0, 0.2, 1, 2, 5, 10 Hz daily for 3 days. Mean illumination was 680 lux. Changes in ocular components and corneal curvature were measured. RESULTS Refraction, eye length, and choroidal changes were dependent on the presence of blue light (P < 0.03, all) and on temporal frequency (P < 0.03, all). In the presence of blue light, refraction did not change across frequencies (mean change -0.24 [diopters] D), while in the absence of blue light, we observed a hyperopic shift (>1 D) at high frequencies, and a myopic shift (>-0.6 D) at low frequencies. With blue light there was little difference in eye growth across frequencies (77 μm), while in the absence of blue light, eyes grew more at low temporal frequencies and less at high temporal frequencies (10 vs. 0.2 Hz: 145 μm; P < 0.003). Overall, neonatal astigmatism was reduced with blue light. CONCLUSIONS Illuminants rich in blue light can protect against myopic eye growth when the eye is exposed to slow changes in luminance contrast as might occur with near work.

Potential signal to accommodation from the Stiles–Crawford effect and ocular monochromatic aberrations

The purpose of this study is to determine if cues within the blurred retinal image due to the Stiles–Crawford (SC) effect and the eyes monochromatic aberrations can drive accommodation with a small pupil (3 mm) that is typical of bright photopic conditions. The foveal, psychophysical SC function (17 min arc) and ocular monochromatic aberrations were measured in 21 visually normal adults. The retinal image of a 10.2 min arc disc was simulated for spherical defocus levels of −1 D, 0 D and +1 D in each of four conditions consisting of combinations of the presence or absence of the individual SC function and monochromatic aberrations with a 3 mm pupil. Accommodation was recorded in 11 participants as each viewed the simulations through a 0.75 mm pinhole. The SC effect alone did not provide a significant cue to accommodation. Monochromatic aberrations provided a statistically significant but rather small cue to monocular accommodation.

The role of temporal contrast and blue light in emmetropization

HighlightsThe eye is vulnerable to myopia with exposure to low contrast visual stimuli.Exposure to low temporal frequency visual stimuli increases vulnerability.As does a yellowish light source without a strong blue component. Abstract A previous experiment showed that blue light (as a component of white light) protected against low temporal frequency dependent eye growth. This experiment investigated the role of temporal contrast. White leghorn chicks were exposed to white (with blue) or yellow (without blue) LED lighting modulated at either low (0.2 Hz) or high (10 Hz) temporal frequencies. Four cone contrast conditions were used: low (16%), medium (32%), medium–high (60%) and very‐high (80%). Chicks were exposed to the lighting condition for 3 days (mean 680 lux). Exposure to high temporal frequencies, with very high temporal contrast, reduced eye growth, regardless of spectral content. However, at low temporal frequencies, eye growth was dependent on the illuminant. At lower temporal contrast levels, growth increased regardless of temporal or spectral characteristics. To conclude, very high temporal contrast, white light, provides a “stop” signal for eye growth that overrides temporal cues for growth that manifest in yellow light.