Temporal color contrast guides emmetropization in chick.

As a result of longitudinal chromatic aberration (LCA), longer wavelengths are blurred when shorter wavelengths are in focus, and vice versa. As a result, LCA affects the color and temporal aspects of the retinal image with hyperopic defocus. In this experiment, we investigated how the sensitivity to temporal color contrast affects emmetropization. Ten-day-old chicks were exposed for three days to sinusoidal color modulation. The modulation was either blue/yellow flicker (BY) (n = 57) or red/green flicker (RG) (n = 60) simulating hyperopic defocus with and without a blue light component. The color contrasts tested were 0.1, 0.2, 0.3, 0.4, 0.6, and 0.8 Michelson contrast. The mean illuminance of all stimuli was 680 lux. Temporal modulation was either of a high (10 Hz) or low (0.2 Hz) temporal frequency. To test the role of short- and double-cone stimulation, an additional condition silenced these cones in RG_0.4 (D-) and was compared with RG_0.4 (D+) (n = 14). Changes in ocular components and refractive error were measured using Lenstar and a photorefractometer. With high temporal frequency BY representing an in-focus condition for shorter-wavelengths, we found that high temporal frequency BY contrast was positively correlated with vitreous expansion (R2 = 0.87, p < 0.01), expanding the vitreous to compensate for hyperopic defocus. This expansion was offset by low temporal frequency RG, which represented blurred longer wavelengths. The reduction in vitreous expansion in RG_0.4, was enhanced in D+ compared to D- (p < 0.001), indicating a role for short- and/or double-cones. With high temporal frequency RG representing an in-focus condition for longer-wavelengths, we found that high temporal frequency RG contrast was also positively correlated with a linear increase in vitreous chamber depth (R2 = 0.84, p < 0.01) and eye length (R2 = 0.30, p ≤ 0.05), required to compensate for hyperopic defocus, but also with RG sensitive choroidal thickening (R2 = 0.18: p < 0.0001). These increases in the vitreous and eye length were enhanced with D+ compared to D- (p = 0.003) showing the role of short- and double-cones in finessing the vitreous response to hyperopic defocus. Overall, the increase in vitreous chamber depth in RG was offset by reduced expansion in BY, indicating sensitivity to the shorter focal length of blue light and wavelength defocus. Predictable changes in cone contrast and temporal frequency of the retinal image that occur with LCA and defocus result in homeostatic control of emmetropization.

Signals for defocus arise from longitudinal chromatic aberration in chick.

Chicks respond to two signals from longitudinal chromatic aberration (LCA): a wavelength defocus signal and a chromatic signal. Wavelength defocus predicts reduced axial eye growth in monochromatic short-wavelength light, compared to monochromatic long-wavelength light. Wavelength defocus may also influence growth in broadband light. In contrast, a chromatic signal predicts increased growth when short-wavelength contrast > long-wavelength contrast, but only when light is broadband. We aimed to investigate the influence of blue light, temporal frequency and contrast on these signals under broadband conditions. Starting at 12 to 13 days-old, 587 chicks were exposed to the experimental illumination conditions for three days for 8h/day and spent the remainder of their day in the dark. The stimuli were flickering lights, with a temporal frequency of 0.2 or 10 Hz, low (30%) or high contrast (80%), and a variety of ratios of cone contrast simulating the effects of defocus with LCA. There were two color conditions, with blue contrast (bPlus) and without (bMinus). Stimuli in the "bPlus" condition varied the amounts of long- (L), middle- (M_) and double (D-) cone contrast, relative to short- (S-) and (UV-) cone contrast, to simulate defocus. Stimuli in the "bMinus" condition only varied the relative modulations of the L + D vs. M cones. In all cases, the average of the stimuli was white, with an illuminance of 777 lux, with cone contrast created through temporal modulation. A Lenstar LS 900 and a Hartinger refractometer were used to measure ocular components and refraction. Wavelength defocus signals with relatively high S-cone contrast resulted in reduced axial growth, and more hyperopic refractions, under low-frequency conditions (p = 0.002), in response to the myopic defocus of blue light. Chromatic signals with relatively high S-cone contrast resulted in increased axial growth and more myopic refractions, under high frequency, low contrast, conditions (p < 0.001). We conclude that the chromatic signals from LCA are dependent on the temporal frequency, phase, and relative contrast of S-cone temporal modulation, and recommend broadband spectral and temporal environments, such as the outdoor environment, to optimize the signals-for-defocus in chick.

The role of dopamine in eye growth responses to color and luminance flicker in chicks.

Stroboscopic luminance flicker has been found to prevent the increase in eye growth normally associated with form deprivation through the release of retinal dopamine. In this study, we sought to investigate whether dopamine plays a role in the decreased growth observed with 2 Hz sine-wave luminance flicker and increased growth with color flicker. Starting 5-7 days after hatching, chicks were exposed to 2 Hz sinusoidally modulated illumination (Mean: 680 lux) for 4 days and were otherwise in the dark. Chicks were exposed to color-modulated red and green (RG) light, to luminance modulated RGB components (LUM), or to a no-flicker (NF) control. Chicks received daily 10 µL intravitreal injections of apomorphine, spiperone, or saline. Fellow eyes received no injection. Spiperone injections prevented the decrease in eye growth typically seen with LUM flicker, with a relative increase in eye length, but no other significant effects compared with saline controls. Apomorphine injections prevented the increase in eye growth typically seen with RG flicker, with a relative decrease in eye length compared to saline controls. These results indicate a role for the activation of D2-receptor types in the inhibition of eye growth in response to luminance flicker, and a lack of dopamine receptor activation associated with the increase in eye growth with color flicker.

The role of luminance and chromatic cues in emmetropisation.

PURPOSE: 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. RECENT FINDINGS: There are three ways in which changes in luminance and colour contrast could provide cues: (1) The eye could maximize luminance contrast. Monochromatic light experiments have shown that the human eye can accommodate and animal eyes can emmetropise using changes in luminance contrast alone. However, by reducing the effectiveness of luminance cues in monochromatic and white light by introducing astigmatism, or by reducing light intensity, investigators have revealed that the eye also uses colour cues in emmetropisation. (2) The eye could compare relative cone contrast to derive the sign of defocus information from colour cues. Experiments involving simulations of the retinal image with defocus have shown that relative cone contrast can provide colour cues for defocus in accommodation and emmetropisation. In the myopic simulation the contrast of the red component of a sinusoidal grating was higher than that of the green and blue component and this caused relaxation of accommodation and reduced eye growth. In the hyperopic simulation the contrast of the blue component was higher than that of the green and red components and this caused increased accommodation and increased eye growth. (3) The eye could compare the change in luminance and colour contrast as the eye changes focus. An experiment has shown that changes in colour or luminance contrast can provide cues for defocus in emmetropisation. When the eye is exposed to colour flicker the eye grows almost twice as much, and becomes more myopic, compared to when the eye is exposed to luminance flicker. SUMMARY: Neural responses of the luminance and colour mechanisms direct accommodation and emmetropisation mechanisms to different focal planes. Therefore, it is likely that the set point of refraction and accommodation is dependent on the sensitivity of the eye to changes in spatial and temporal, colour and luminance contrast.

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.

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.

Sensitivity to S-Cone Stimuli and the Development of Myopia.

Purpose: Longitudinal chromatic aberration (LCA) is a color signal available to the emmetropization process that causes greater myopic defocus of short wavelengths than long wavelengths. We measured individual differences in chromatic sensitivity to explore the role LCA may play in the development of refractive error. Methods: Forty-four observers were tested psychophysically after passing color screening tests and a questionnaire for visual defects. Refraction was measured and only subjects with myopia or hyperopia without severe astigmatism participated. Psychophysical detection thresholds for 3 cyc/deg achromatic, L-, M-, and S-cone-isolating Gabor patches and low-frequency S-cone increment (S+) and decrement (S-) blobs were measured. Parametric Pearson correlations for refractive error versus threshold were calculated and nonparametric bootstrap 95% percentage confidence intervals (BCIs) for r were computed. Results: S-cone Gabor detection thresholds were higher than achromatic, L-, and M-cone Gabors. S-cone Gabor thresholds were higher than either S+ or S- blobs. These results are consistent with studies using smaller samples of practiced observers. None of the thresholds for the Gabor stimuli were correlated with refractive error (RE). A negative correlation with RE was observed for both S+ (r = -0.28; P = 0.06; BCI: r = -0.5, -0.04) and S- (r = -0.23; P = 0.13; BCI = -0.46, 0.01) blobs, although this relationship did not reach conventional statistical significance. Conclusions: Thresholds for S+ and S- stimuli were negatively related to RE, indicating that myopes may have reduced sensitivity to low spatial frequency S-cone stimuli. This reduced S-cone sensitivity might have played a role in their failure to emmetropize normally.

Effect of third-order aberrations on dynamic accommodation.

We investigate the potential for the third-order aberrations coma and trefoil to provide a signed cue to accommodation. It is first demonstrated theoretically (with some assumptions) that the point spread function is insensitive to the sign of spherical defocus in the presence of odd-order aberrations. In an experimental investigation, the accommodation response to a sinusoidal change in vergence (1-3D, 0.2Hz) of a monochromatic stimulus was obtained with a dynamic infrared optometer. Measurements were obtained in 10 young visually normal individuals with and without custom contact lenses that induced low and high values of r.m.s. trefoil (0.25, 1.03 microm) and coma (0.34, 0.94 microm). Despite variation between subjects, we did not find any statistically significant increase or decrease in the accommodative gain for low levels of trefoil and coma, although effects approached or reached significance for the high levels of trefoil and coma. Theoretical and experimental results indicate that the presence of Zernike third-order aberrations on the eye does not seem to play a crucial role in the dynamics of the accommodation response.

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 individual's 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.

Opposing effects of atropine and timolol on the color and luminance emmetropization mechanisms in chicks.

This study analyzed the luminance and color emmetropization response in chicks treated with the nonselective parasympathetic antagonist atropine and the sympathetic ß-receptor blocker timolol. Chicks were binocularly exposed (8h/day) for 4days to one of three illumination conditions: 2Hz sinusoidal luminance flicker, 2Hz sinusoidal blue/yellow color flicker, or steady light (mean 680lux). Atropine experiments involved monocular daily injections of either 20µl of atropine (18nmol) or 20µl of phosphate-buffered saline. Timolol experiments involved monocular daily applications of 2 drops of 0.5% timolol or 2 drops of distilled H2O. Changes in the experimental eye were compared with those in the fellow eye after correction for the effects of saline/water treatments. Atropine caused a reduction in axial length with both luminance flicker (-0.078±0.021mm) and color flicker (-0.054±0.017mm), and a reduction in vitreous chamber depth with luminance flicker (-0.095±0.023mm), evoking a hyperopic shift in refraction (3.40±1.77D). Timolol produced an increase in axial length with luminance flicker (0.045±0.030mm) and a myopic shift in refraction (-4.07±0.92D), while color flicker caused a significant decrease in axial length (-0.046±0.017mm) that was associated with choroidal thinning (-0.046±0.015mm). The opposing effects on growth and refraction seen with atropine and timolol suggest a balancing mechanism between the parasympathetic and ß-receptor mediated sympathetic system through stimulation of the retina with luminance and color contrast.

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.

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.

The role of short-wavelength sensitive cones and chromatic aberration in the response to stationary and step accommodation stimuli.

The aim of the experiment was to test for a contribution from short-wavelength sensitive cones to the static and step accommodation response, to compare responses from short and long- plus middle-wavelength sensitive cone types, and to examine the contribution of a signal from longitudinal chromatic aberration to the accommodation response. Accommodation was monitored continuously (eight subjects) to a square-wave grating (2.2 c/d; 0.57 contrast) in a Badal optometer. The grating stepped (1.00 D) randomly towards or away from the eye from a starting position of 2.00 D. Five illumination conditions were used to isolate cone responses, and combine them with or without longitudinal chromatic aberration. Accuracy of the response before the step, step amplitude, latencies and time-constants, were compared between conditions using single factor ANOVA and t-test comparisons. Both S-cones and LM-cones mediated static and step accommodation responses. S-cone contrast drives "static" accommodation for near, but the S-cone response is too slow to influence step dynamics when LM-cones participate.

Accommodation to simulations of defocus and chromatic aberration in the presence of chromatic misalignment.

Previous studies have demonstrated that accommodation will respond to sine gratings in which the relative modulations of red, green and blue image components have been altered to simulate the effects of defocus and longitudinal chromatic aberration. The present study aimed to determine the tolerance of the accommodative system to relative phase shifts in those components induced by chromatic misalignment. It was found that accommodation can tolerate moderate amounts of chromatic misalignment (6'), but responds adversely when misalignments are large. Applications to visual display terminals and spectacle lens and instrument design are discussed.

Chick eyes compensate for chromatic simulations of hyperopic and myopic defocus: evidence that the eye uses longitudinal chromatic aberration to guide eye-growth.

Longitudinal chromatic aberration (LCA) causes short wavelengths to be focused in front of long wavelengths. This chromatic signal is evidently used to guide ocular accommodation. We asked whether chick eyes exposed to static gratings simulating the chromatic effects of myopic or hyperopic defocus would "compensate" for the simulated defocus. We alternately exposed one eye of each chick to a sine-wave grating (5 or 2 cycle/deg) simulating myopic defocus ("MY defocus": image focused in front of retina; hence, red contrast higher than blue) and the other eye to a grating of the same spatial frequency simulating hyperopic defocus ("HY defocus": blue contrast higher than red). The chicks were placed in a drum with one eye covered with one grating, and then switched to another drum with the other grating with the other eye covered. To minimize the effects of altered eye-growth on image contrast, we studied only the earliest responses: first, we measured changes in choroidal thickness 45 min to 1 h after one 15-min episode in the drum, then we measured glycosaminoglycans (GAG) synthesis in sclera and choroid (by the incorporation of labeled sulfate in tissue culture) after a day of four 30-min episodes in the drum. The eyes compensated in the appropriate directions: The choroids of the eyes exposed to the HY simulation showed significantly more thinning (less thickening) over the course of the experiment than the choroids of the eyes exposed to the MY simulation (all groups mean:-17 microm; 5 c/d groups: -24 microm; paired t-test (one-tailed): p=0.0006). The rate of scleral GAG synthesis in the eye exposed to the HY simulation was significantly greater than in the eye exposed to the MY simulation (HY/MY ratio=1.20; one sample t-test (one-tailed): p=0.015). There was no significant interaction between the sign of the simulated defocus and either the spatial frequency or the presence of a +3 D lens used to compensate for the 33 cm distance of the drum. Although previous work has shown that chromatic cues to defocus are not essential for lens-compensation, in that chicks can compensate in monochromatic light, our evidence implies that the eye may be able to infer whether the eye is myopic or hyperopic from the different chromatic contrasts that result from different signs of defocus.

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 eye's 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 eleven 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.